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					                                                Cyto-nuclear genomic dissociation in African elephant species
                                                                                     Alfred L. Roca*, Nicholas Georgiadis‡ and Stephen J. O’Brien†
                                                *Basic   Research Program, SAIC-Frederick, and †Laboratory of Genomic Diversity, National Cancer Institute, Frederick, MD 21702, USA
                                                                                       ‡Mpala Research Center, PO Box 555, Nanyuki, Kenya




FIGURE 1: INTRODUCTION                                                                                                                                                                                                                                                     (A) mtDNA ND5
                                                                                                                                                                                                                                                                               319 bp
                                                                                                                                                                                                                                                                                                                                                                                                   (B) Y chromosome AMELY
                                                                                                                                                                                                                                                                                                                                                                                                   1551 bp

Multiple lines of evidence suggest that forest and savanna elephants are distinct species.                                                                                     FIGURE 3                                                                                                                      DS (5)



Measurements from 295 skulls demonstrated that forest and savanna elephants fall into two morphologically                                                                                                                                                                                                DS (11)
                                                                                                                                                                                                                                                                                                         BE (1)                                                                                                                                                    BE4059
                                                                                                                                                                                                                                                                                                                                                                                                                                                                            DS1537
                                                                                                                                                                                                                                                                                                                                                                                                                                                                            DS1556
                                                                                                                                                                                                                                                                                                                                                                                                                                                                            LO3517




distinct groups (left panel; Groves and Grubb, 2000). Nuclear gene analyses using both slower-evolving                                                                         Maternally       and       paternally                                                                                     GR (1)                                                                                                                                    63/80/64        DS1504
                                                                                                                                                                                                                                                                                                                                                                                                                                                                   DS1521
                                                                                                                                                                                                                                                                                                                                                                                                                                                                   DS1523
                                                                                                                                                                                                                                                                                                                                                                                                                                                                   DS1524
                                                                                                                                                                                                                                                                                                                                                                                                                                                                   DS1555
                                                                                                                                                                                                                                                                                                                                                                                                                                                                   OD0001




nuclear gene sequences (center; Roca et al., 2001) and more rapidly evolving microsatellites (right; Comstock                                                                  inherited markers demonstrate                                                                                                 DS (8)
                                                                                                                                                                                                                                                                                                                                                                                                                                   99/99/98
                                                                                                                                                                                                                                                                                                                                                                                                                                                         DS1503
                                                                                                                                                                                                                                                                                                                                                                                                                                                         DS1505
                                                                                                                                                                                                                                                                                                                                                                                                                                                         DS1511
                                                                                                                                                                                                                                                                                                                                                                                                                                                         DS1527
                                                                                                                                                                                                                                                                                                                                                                                                                                                                   GR0016
                                                                                                                                                                                                                                                                                                                                                                                                                                                                   GR0022




                                                                                                                                                                               cytonuclear             dissociation.
                                                                                                                                                                                                                                                                                                                                                                                                                                                         DS1528
                                                                                                                                                                                                                                                                                                                                                                                                                                                         DS1530
                                                                                                                                                                                                                                                                                                                                                                                                                                              II
et al., 2002) demonstrated a deep genetic split between forest and savanna elephants, estimated at 3.5 million
                                                                                                                                                                                                                                                                                                                                                                                                                                                         DS1532
                                                                                                                                                                                                                                                                                                                                                                                                                                                         DS1543
                                                                                                                                                                                                                                                                                                                                                                                                                                                         LO3505

                                                                                                                                                                                                                                                                           62/74/78                          DS (1), OD (2)            BE (2), WA (1)                                                                     AM0003
                                                                                                                                                                                                                                                                                                                                                                                                                          AM0005



                                                                                                                                                                               Phylogenetic relationships for Asian,
                                                                                                                                                                                                                                                                                                                                                                                                                          AM0007
                                                                                                                                                                                                                                                                                                                                                                                                                          AM0015



years. Only a few morphological intermediates and genetic hybrids were detected in a zone of mixed habitat
                                                                                                                                                                                                                                                                                                                                                                                                                          AM0018
                                                                                                                                                                                                                                                                                      IIB                                                                                                                                 AM0019
                                                                                                                                                                                                                                                                                                                                                                                                                          AM0020
                                                                                                                                                                                                                                                                                                                                                                                                                          AM0021
                                                                                                                                                                                                                                                                                                          CH (1), HW (6), SA (8), ZZ (4)                                                                                  AM0023




that surrounds the tropical forests of Africa.                                                                                                                                 African forest, and African savanna                                                                                 HW (2), NG (4), SA (2), SE (9), TA (1), ZZ (1)
                                                                                                                                                                                                                                                                                                                                                                                                                          AM0030
                                                                                                                                                                                                                                                                                                                                                                                                                          AM0035
                                                                                                                                                                                                                                                                                                                                                                                                                          AM4551
                                                                                                                                                                                                                                                                                                                                                                                                                          AM4576
                                                                                                                                                                                                                                                                                                                                                                                                                          AM4583
                                                                                                                                                                                                                                                                                                                                                                                                                          AM4584
                                                                                                                                                                                                                                                                                                                                                                                                                          AM4585




   In contrast to the distinctions between forest and savanna elephants detected by morphological and nuclear                                                                  elephants inferred from (A) 319 bp of                                                 89/92/97                      GR (10)
                                                                                                                                                                                                                                                                                                                                                                                                                          AM4587
                                                                                                                                                                                                                                                                                                                                                                                                                          BE4035
                                                                                                                                                                                                                                                                                                                                                                                                                          BE4053
                                                                                                                                                                                                                                                                                                                                                                                                                          CH0882
                                                                                                                                                                                                                                                                                                                                                                                                                          CH0885
                                                                                                                                                                                                                                                                                                                                                                                                                          CH0895
                                                                                                                                                                                                                                                                                                                                                                                                                          CH0931




genetic studies, analyses using mitochondrial DNA (mtDNA) have detected genetic diversity in savanna                                                                           the        maternally        inherited                                                           II                                                                                                                                        HW0062
                                                                                                                                                                                                                                                                                                                                                                                                                          HW0067
                                                                                                                                                                                                                                                                                                                                                                                                                          HW0076
                                                                                                                                                                                                                                                                                                                                                                                                                          HW0082
                                                                                                                                                                                                                                                                                                                                                                                                                          HW0086
                                                                                                                                                                                                                                                                                                                                                                                                                          HW0092
                                                                                                                                                                                                                                                                                                                                                                                                                          HW0112




elephants high enough to appear incongruent with nuclear DNA studies, and suggested greater mixing                                                                             mitochondrial ND5 gene (number of                                                                                              DS (1), LO (9), OD (1)

                                                                                                                                                                                                                                                                                                              BE (1), WA (1)
                                                                                                                                                                                                                                                                                                                                                                                                                          HW0115
                                                                                                                                                                                                                                                                                                                                                                                                                          HW0117
                                                                                                                                                                                                                                                                                                                                                                                                                          HW0120
                                                                                                                                                                                                                                                                                                                                                                                                                          HW0122
                                                                                                                                                                                                                                                                                                                                                                                                                          HW0124
                                                                                                                                                                                                                                                                                                                                                                                                                          HW0151
                                                                                                                                                                                                                                                                                                                                                                                                                          KE4501




between forest and savanna elephants. To investigate this apparent disparity, we sampled wild elephant tissue                                                                  individuals with identical haplotypes                                                                  94/96/98

                                                                                                                                                                                                                                                                                                   IIA
                                                                                                                                                                                                                                                                                                                    LO (2)
                                                                                                                                                                                                                                                                                                                                                                                                                          KE4509
                                                                                                                                                                                                                                                                                                                                                                                                                          KE4511
                                                                                                                                                                                                                                                                                                                                                                                                                          KE4516
                                                                                                                                                                                                                                                                                                                                                                                                                          KE4517
                                                                                                                                                                                                                                                                                                                                                                                                                          KE4549
                                                                                                                                                                                                                                                                                                                                                                                                                          KE4550
                                                                                                                                                                                                                                                                                                                                                                                                                          KE4601




from 21 African locations to determine the DNA sequences for 1642 biparentally inherited chromosomal                                                                           indicated by location), and from (B)                                                                                                BE (1), WA (2)
                                                                                                                                                                                                                                                                                                                                                                                                                          KE4607
                                                                                                                                                                                                                                                                                                                                                                                                                          KE4609
                                                                                                                                                                                                                                                                                                                                                                                                                          KE4614
                                                                                                                                                                                                                                                                                                                                                                                                                          KE4617
                                                                                                                                                                                                                                                                                                                                                                                                                          KE4620
                                                                                                                                                                                                                                                                                                                                                                                                                          KE4621
                                                                                                                                                                                                                                                                                                                                                                                                                          KE4623




segments in 3 X-linked genes, and 302 mtDNA and 128 Y chromosome sequences.                                                                                                    1551 bp of the paternally inherited Y                                                                                                                                                                                                      KR0114
                                                                                                                                                                                                                                                                                                                                                                                                                          MA0807
                                                                                                                                                                                                                                                                                                                                                                                                                          NA4653
                                                                                                                                                                                                                                                                                                                                                                                                                          NA4660
                                                                                                                                                                                                                                                                                                                                                                                                                          NA4665
                                                                                                                                                                                                                                                                                                                                                                                                                          NA4668
                                                                                                                                                                                                                                                                                                                                                                                                                          NA4669



                                                                                                                                                                               chromosome gene AMELY (each                                                                                                            90/91/87
                                                                                                                                                                                                                                                                                                                                           AB (6), AM (34), CH (5), KE (16), KR (12) MA (1),
                                                                                                                                                                                                                                                                                                                                           MK (1), NA (21), NG (3), SE (1), SW (5), TA (6)
                                                                                                                                                                                                                                                                                                                                                                                                                          NA4670
                                                                                                                                                                                                                                                                                                                                                                                                                          NA4671
                                                                                                                                                                                                                                                                                                                                                                                                                          NA4675
                                                                                                                                                                                                                                                                                                                                                                                                                          NA4678
                                                                                                                                                                                                                                                                                                                                                                                                                          NA4688
                                                                                                                                                                                                                                                                                                                                                                                                                          NA4697
                                                                                                                                                                                                                                                                                                                                                                                                                          NA4699



                                                                                                                                                                               individual shown separately). Forest                                                                                         70/66/63                           AB (1), AM (2), KE (4)
                                                                                                                                                                                                                                                                                                                                                                                                       87/93/90

                                                                                                                                                                                                                                                                                                                                                                                                                    I
                                                                                                                                                                                                                                                                                                                                                                                                                          NA4702
                                                                                                                                                                                                                                                                                                                                                                                                                          NA4704
                                                                                                                                                                                                                                                                                                                                                                                                                          NA4710
                                                                                                                                                                                                                                                                                                                                                                                                                          NG2178
                                                                                                                                                                                                                                                                                                                                                                                                                          NG2180
                                                                                                                                                                                                                                                                                                                                                                                                                          NG2181
                                                                                                                                                                                                                                                                                                                                                                                                                          NG2182



                                                                                                                                                                               populations or individuals are                                                                                                                     CH (3), HW (4), KR (25), MA (3), SW (5)
                                                                                                                                                                                                                                                                                                                                                                                                                          NG2191
                                                                                                                                                                                                                                                                                                                                                                                                                          NG2192
                                                                                                                                                                                                                                                                                                                                                                                                                          NG2193
                                                                                                                                                                                                                                                                                                                                                                                                                          NG2194
                                                                                                                                                                                                                                                                                                                                                                                                                          NG2214
                                                                                                                                                                                                                                                                                                                                                                                                                          NG2215
                                                                                                                                                                                                                                                                                                                                                                                                                          NG2229



                                                                                                                                                                               indicated in green; savanna in blue;                                                                  100/100/100
                                                                                                                                                                                                                                                                                                                                                                                                                          SA0972
                                                                                                                                                                                                                                                                                                                                                                                                                          SA0993
                                                                                                                                                                                                                                                                                                                                                                                                                          SA0994
                                                                                                                                                                                                                                                                                                                                                                                                                          SA1002
                                                                                                                                                                                                                                                                                                                                                                                                                          SA1004
                                                                                                                                                                                                                                                                                                                      I                                                                                                   SA1005
                                                                                                                                                                                                                                                                                                                                                                                                                          SA1009



                                                                                                                                                                               Asian elephants in red. Garamba                                                                                                                   AB (1), KE (17), WA (4)
                                                                                                                                                                                                                                                                                                                                                                                                                          SE2051
                                                                                                                                                                                                                                                                                                                                                                                                                          SE2098
                                                                                                                                                                                                                                                                                                                                                                                                                          SE2101
                                                                                                                                                                                                                                                                                                                                                                                                                          SE2103
                                                                                                                                                                                                                                                                                                                                                                                                                          SE2104
                                                                                                                                                                                                                                                                                                                                                                                                                          SE2106
                                                                                                                                                                                                                                                                                                         64/64/63                                                                                                         SE2165



                                                                                                                                                                               (GR) is a mixed habitat zone and is                                                                                                               GR (1)                                                                                   TA1144
                                                                                                                                                                                                                                                                                                                                                                                                                          TA1145
                                                                                                                                                                                                                                                                                                                                                                                                                          TA1431
                                                                                                                                                                                                                                                                                                                                                                                                                          TA1443
                                                                                                                                                                                                                                                                                                                                                                                                                          TA1450
                                                                                                                                                                                                                                                                                                                                                                                                                          TA1458
                                                                                                                                                                                                                                                                                                                                     BE (1), WA (1)

                                                                                                                                                                               not colored.
                                                                                                                                                                                                                                                                                                                                                                                                                          WA4020
                                                                                                                                                                                                                                                                                                                                                                                                                          WA4022
                                                                                                                                                                                                                                                                                                                                                                                                                          WA4027
                                                                                                                                                                                                                                                                                                                                                                                                                          ZZ0145
                                                                                                                                                                                                                                                                                                                                                                                                                          ZZ0148
                                                                                                                                                                                                                                                                                                                                                                                                                                                                            Ema006



                                                                                                                                                                                                                                                                                                                                               Elephas maximus (1)                                                0.001 substitutions/site
                                                                                                                                                                                                                                                                           0.005 substitutions/site




                                                                                                                                                                                     mtDNA biparental Y chro.

                                                                                                                                                                                                                                                                                 AB
                                                                                                                                                                                                                                                                                      mtDNA biparental Y chro.                            FIGURE 4
                                                                                                                                                                                LO                                           WA                                                  KE
                                                                                                                                                                                                                                                                                 MK
                                                                                                                                                                                         n=11          n=65        n=2
                                                                                                                                                                                                                             BE                                                               n=58        n=280           n=15            Distribution of cytonuclear disequilibrium.
                                                                                                                                                                                OD
                                                                                                                                                                                             3            7         1
                                                                                                                                                                                                                                                                                 AM                                                       Pie charts indicate by locale the distribution of
                                                                                                                                                                                                                                               R                                              35           149            17

                                                                                                                                                                                DS                                                                        KE
                                                                                                                                                                                                                                                             MK
                                                                                                                                                                                                                                                                                 CH
                                                                                                                                                                                                                                                                                          8
                                                                                                                                                                                                                                                                                                                                          genetic markers that are inherited maternally
                                                                                                                                                                                                                              OD                          AB                     SA
                                                                                                                                                                                     1
                                                                                                                                                                                             26
                                                                                                                                                                                                      3
                                                                                                                                                                                                          171      15
                                                                                                                                                                                                                                                        SE AM                          4
                                                                                                                                                                                                                                                                                              11           154            11
                                                                                                                                                                                                                                                                                                                                          (left pie chart in each set of three), paternally
                                                                                                                                                                                GR                                                                      NG TA
                                                                                                                                                                                             11           48
                                                                                                                                                                                                               1
                                                                                                                                                                                                                    3
                                                                                                                                                                                                                                                                                HW
                                                                                                                                                                                                                                                                                               8            71            13
                                                                                                                                                                                                                                                                                                                                          (right), or biparentally (center). Totals indicate
                                                                                                                                                                                         1                               1
                                                                                                                                                                                BE                                                                                                   SW
                                                                                                                                                                                                                                                                                     ZZ
                                                                                                                                                                                                                                                                                                     5
                                                                                                                                                                                                                                                                                                                                          the number of individuals (mtDNA, Y
                                                                                    Savannah elephants   Savannah elephants   Savannah elephants        Forest elephants
                                                                                    SOUTHERN             EASTERN              NORTH-CENTRAL             DS-Dzanga Sangha
                                                                                                                                                                                             5
                                                                                                                                                                                                  4
                                                                                                                                                                                                       26
                                                                                                                                                                                                               1
                                                                                                                                                                                                                    2
                                                                                                                                                                                                                                                                                              10            46             2
                                                                                                                                                                                                                                                                                                                                          chromosomes) or combined number of
                                                                                    CH-Chobe             AB-Aberdares         BE-Benoue                 LO-Lope                 WA                                                                                              MA
                                                                                    HW-Hwange
                                                                                    KR-Kruger
                                                                                                         AM-Amboseli
                                                                                                         MK-Mount Kenya
                                                                                                                              WA-Waza                   GA-Garamba
                                                                                                                                                                                         1
                                                                                                                                                                                             5            94        3                      ZZ SW                                               4            56             1
                                                                                                                                                                                                                                                                                                                                          chromosome segments (biparental genes)
                                                                                    NA-Namibia           KE-Central Kenya                                                                                                         NA     CH HW
                                                                                                                                                   16 microsatellite loci
                                                                                    MA-Mashatu
                                                                                    SA-Savuti
                                                                                                         NG-Ngorongoro
                                                                                                         SE-Serengeti
                                                                                                                              Asian elephants
                                                                                                                                                                                SE                                                     SA
                                                                                                                                                                                                                                                                                 KR                                                       examined.
                                                                                    SW-Sengwa            TA-Tarangire                              Comstock et al. 2002                                                                      MA
                                                                                    ZZ-Zambezi                                                                                       3
                                                                                                                                                                                             9            44        7
                                                                                                                                                                                                                                                   KR                                         37           170             2
                                                                                                                                                                                                                                                                                                                                              Map indicates locations of sampled elephant
                                                                                                                                                                                NG                                                                                               NA
                                                                                                                                                                                             4
                                                                                                                                                                                                  1
                                                                                                                                                                                                          32       11
                                                                                                                                                                                                                                                                                              21           157            15
                                                                                                                                                                                                                                                                                                                                          populations in Africa. Green circles are forest
                                                                                                                                                                                TA                                                                                                     ND5                BGN
                                                                                                                                                                                                                                                                                                          PLP
                                                                                                                                                                                                                                                                                                               AMELY                      locations. Blue circles are savanna locations.
                                                                                                                                                                                             6            67        7

                                                                                                                                                                                     ND5               BGN  AMELY
                                                                                                                                                                                                                                                                                                         PHKA2
                                                                                                                                                                                                                                                                                                                                          Garamba (GR) includes both habitats. Orange
                                                                                                                                                                                                       PLP
                                                                                                                                                                                                      PHKA2                                                                                                                               indicates current African elephant range; historic
                                                                                                                                                                                DS-Dzanga Sangha, LO-Lope, OD-Odzala, GR-Garamba, AB-Aberdares, AM-                                                                                       range includes entire land area shown.
                                                                                                                                                                                Amboseli, BE-Benoue, CH-Chobe, HW-Hwange, KE-Central Kenya, KR-Kruger,
                                                                                                                                                                                MA-Mashatu, MK-Mount Kenya, NA-Namibia, NG-Ngorongoro, SA-Savuti, SE-
METHODS                                                                                                                                                                         Serengeti, SW-Sengwa, TA-Tarangire, WA-Waza, ZZ-Zambezi.


DNA was extracted from samples from wild African elephants and captive Asian elephants (Elephas maximus). Three nuclear
gene segments (BGN, PHKA2 and PLP); a portion of the mitochondrial gene ND5, and a Y-chromosome gene fragment (AMELY)
were amplified and sequenced. Sequences were aligned using CLUSTALX. Phylogenetic analyses were performed using
                                                                                                                                                                              FIGURE 5: CONCLUSIONS
maximum parsimony (MP), neighbor joining (NJ), and maximum likelihood (ML) methods implemented in PAUP*4.0b10.
                                                                                                                                                                              Cytonuclear disequilibrium suggests historic unidirectional hybridization (i.e., savanna males and forest
                                                                                                                                                                              females) with subsequent unidirectional backcrossing to larger reproductively successful savanna males,
     African
                                                                                                                                                                              swamping the forest nuclear genomic contribution. The interactions between forest and savanna elephants
                     (A)                                    (B)                                                   (C)
     Forest                                                                                                                                                                   inferred from differing patterns detected by maternally-inherited versus paternally- or biparentally-inherited
                                                                                                                                                                              genes are as follows:                                                                                 Herd#2
                                                                                                                                                                                                                                                                                                                                                                                                       A      Herd#1                                                                 A
                                                                                                                                                                             (A) Male-mediated gene flow occurs between adjacent forest elephant                                                                                                                                                                                      B
                                                                                                                                                                                                                                                                                                                                                                                           Forest
      Asian                                                                                                                                                                  herds, and between adjacent savanna elephant herds; however (B)                                                                                                                                               habitat
                                                                                                                                                                             interbreeding between savanna and forest elephants at the contact zone                                                                                                                                                                                                      C


                                                                                                                                                                             between forest and savanna habitats is rare. (C) As forest habitat
                                                                                                                                                                                                                                                                                                                                                                                               D
                                                                                                                                                                             retreats (or when forest herds move into savanna habitats), larger male
      African
     Savanna                                                                                                                                                                 savanna elephants have increased opportunity to hybridize with forest
                                                                                                                                                                             female elephants. However, (D) the smaller forest and hybrid males do
                                                                                                                                                                             not reproduce due either to outbreeding depression or to reproductive
                           BGN, 646 bp, n=556                     PHKA2, 1012 bp, n=440                                 PLP, 479 bp, n=657
                                                                                                                                                                             dominance by larger unhybridized savanna males. (E) After multiple
                                                                                                                                                                             generations of unidirectional hybridization, nuclear genes alleles are
FIGURE 2                                                                                                                                                                     those of savanna elephants, although a forest mitochondrial haplotype is
                                                                                                                                                                                                                                                                                                                                                                                           Savanna
                                                                                                                                                                                                                                                                                                                                                                                           habitat
                                                                                                                                                                             retained in the now-savanna herds.
Haplotypes for three biparentally-inherited nuclear genes display almost complete separation among                                                                                                                                                                                                                                                                                                 E       Forest mtDNA,
                                                                                                                                                                                                                                                                                                                                                                                                           savanna nuclear DNA
                                                                                                                                                                                                                                                                                                                                                                                                                                                              Savanna mtDNA
                                                                                                                                                                                                                                                                                                                                                                                                                                                              and nuclear DNA
three elephant taxa. MP trees are shown. The length of each gene segment and number of chromosomes
                                                                                                                                                                            References and Acknowledgments
examined are indicated for each gene. Number of chromosomes per haplotype is proportional to the size of
                                                                                                                                                                            Comstock, K. E., Georgiadis, N., Pecon-Slattery, J., Roca, A. L., Ostrander, E. A., O'Brien, S. J. and Wasser, S. K. (2002) Patterns of molecular genetic variation among African elephant populations. Mol Ecol.11:2489-98.
circles; differences between alleles are proportional to the distance between circles. Haplotypes/alleles found                                                             Groves, C. P. and Grubb, P. (2000) Do Loxodonta cyclotis and L. africana interbreed? Elephant 2(4):4-7.
                                                                                                                                                                            Roca, A. L., Georgiadis, N., Pecon-Slattery, J. and O'Brien, S. J. (2001) Genetic evidence for two species of elephant in Africa. Science 293(5534):1473-7.
in Asian elephants are red; African forest haplotypes are green; and African savanna haplotypes are blue. (A)                                                               Roca, A. L., Georgiadis, N. and O'Brien, S. J. (2003) Male-driven genomic chimerization of elephant herds in Africa. In preparation.
BGN haplotypes are completely distinct between forest and savanna populations. (B) PHKA2 proved to be the                                                                   We thank R. Ruggiero, W. J. Murphy, E. Eizirik, A. Brandt, M. P. Gough, B. Gough, M. J. Malasky, J. Arthur, R. L. Hill, D. Munroe, S. Cevario, N. J. Crumpler, G. K. Pei, K. M. Helgen. For elephant samples, we thank A.
                                                                                                                                                                            Turkalo, J. M. Fay, R. Weladji, W. Karesh, M. Lindeque, W. Versvelt, K. Hillman Smith, F. Smith, M. Tchamba, S. Gartlan, P. Aarhaug, A. M. Austmyr, Bakari, Jibrila, J. Pelleteret, L. White, M. Habibou, M. W. Beskreo, D.
most diverse nuclear gene segment; the chromosomes examined were completely distinct between forest and                                                                     Pierre, C. Tutin, M. Fernandez, R. Barnes, B. Powell, G. Doungoubé, M. Storey, M. Phillips, B. Mwasaga, A. Mackanga-Missandzou, B. York and A. Baker at the Burnet Park Zoo, and M. Bush at the National Zoological
                                                                                                                                                                            Park. We thank the governments of Botswana, Cameroon, the Central African Republic, Congo (Brazzaville), Congo (Kinshasa), Gabon, Kenya, Namibia, South Africa, Tanzania, and Zimbabwe for permission to collect
savanna populations.(C) PLP haplotypes were distinct between forest and savanna elephants except for one                                                                    samples. Tissues were obtained in full compliance with specific Federal Fish and Wildlife Permits (endangered/threatened species and CITES Permits US 750138 and US 756611 to N.G.). For funding we thank the U. S.
                                                                                                                                                                            Fish and Wildlife Service, National Geographic Society, and European Union (through the Wildlife Conservation Society). This publication has been funded in part with Federal funds from the National Cancer Institute,
haplotype, indicated by the arrow. This common forest elephant haplotype is present in two individuals from                                                                 National Institutes of Health, under Contract No. N01-CO-12400. The content of this publication does not necessarily reflect the views or policies of the Department of Health and Human Services, nor does mention of trade
                                                                                                                                                                            names, commercial products, or organizations imply endorsement by the U.S. Government.
Cameroon.

				
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