Abstract
Variation in mammalian auditory bulla size has been linked to environmental conditions, and has even been claimed to provide a habitat-specific survival value. Enlarged bullae are typically shared among species adapted to living in arid habitats. Previous studies suggest that jirds also exhibit this adaptive enlargement of the bulla. However, such claims are based on the observation on a limited number of specimens, and thus they provide no quantitative support for this hypothesis. Therefore, we tested this hypothesis using a combination of geometric morphometrics and multivariate statistical techniques on a large sample of two jird species that exhibit a wide and partially overlapping geographical (and hence climatic) range, i.e., Merionescrassus (Sundevall, 1842) and M. libycus (Lichtenstein, 1823). A total of 485 intact skulls of populations originating from Africa to the eastern Iranian Plateau were analysed. The covariation between auditory bulla shape and several geoclimatic variables was explored using the two-block partial least-squares method. To some degree, the results seem to support the hypothesis that morphological variation in the auditory bulla is, indeed, correlated with geoclimatic variation. However, this finding cannot be generalised at several levels, hence refuting the hypothesis of convergent correlations between geoclimatic conditions and bulla morphology. Species-specific differences were found at different levels, suggesting that if local responses are present, they may reflect interactions with phylogenetic constraints.
Appendix
Appendix 1 List of specimens examined (excluding TYPE specimens).
Meriones libycus
Smithsonian Natural Museum of Natural History (Washington D.C. USA)
326941, 326942, 326943, 326944, 326969, 326970, 326971, 326973, 328266, 328267, 328269, 369798, 369799, 369801, 369802, 369806, 369808, 369810, 369812, 369814, 369815, 369817, 369818, 369819, 369822, 369824, 369826, 369827, 369828, 369829, 369831, 369832, 369833, 369834, 354682, 326968, 328257, 328258, 328260, 328262, 328264, 328265, 354690, 354691, 354692, 354695, 341256, 341258, 341261, 341262, 341263, 341264, 341265, 341266, 341267, 328224, 328225, 328271, 328272, 328273, 328274, 328275, 329188, 329189, 329190, 329191, 354696, 354661, 350558, 350559, 350572, 354647, 354649, 354650, 354652, 354653, 354664, 354665.
British Museum of Natural History (London, UK)
40.259, 1983.314, 40.257, 25.4.3.43, 25.4.3.23, 10.3.12.4, 25.4.3.25, 5.7.2.3, 25.4.3.20, 47.383, 56.2.29.5, 19.12.10.3, 47.1412, 47.1414.
Zoological Museum of Ferdowsi University of Mashhad(Mashhad, Iran)
M564, M799, M810, M785, 322, 245, M-35, 1021, 30, 1499, 909, 1473, 1504, 303, 208, 474, 1470, 1468, M844, M571, 1506, 1471, 217, M-17, M-35, 157, M-3, 39, M-28, M-36, 205, 212, M-21, 174, M-29, 1469, 24, M-34, M-7, 518, M-37, 176, 563, M-1, 1482, 574, 461, 88, 9, M-572, M-579, M-12, M-808, 4, M-432, M343, 2, M-331, M-188, M-646, M-196, M-11, 850, 11, M-111, M-842, M-573, 7, M-175, M-548, 14, M-694, M-568, M-236, M-251, 840, 577, 106, M-3, M-12.
Musée National d’Histoire Naturelle (Paris, France)
1950.468, 1950.508, 1957.1330, 1957.1338, 1957.1339, 1958.89, 1958.91, 1958.330, 1958.331, 1958.336, 1958.341, 1958342, 1958343, 1958.344, 1958.345, 1958.346, 1958.347, 1958.351, 1958.352, 1958.750, 1957-334, 1957-335, 1957-332, 1957-336.
Royal Belgian Institute of Natural Sciences (Brussels, Belgium)
10186, 10190, 10189, 10187, 10348, 10322, 10192, 10078, 10137, 10283, 10184, 10497, 10185, 10188, 10318, 10319, 10082, 10282, 10194.
Meriones crassus
Smithsonian Natural Museum of Natural History (Washington D.C. USA)
328235, 328236, 350507, 350508, 350509, 350479, 350481, 329156, 329157, 350468, 350470, 350471, 350472, 350475, 350476, 350477, 350478, 354739, 354741, 354743, 369465, 369466, 350500, 350501, 350503, 350505, 326904, 326905, 326906, 326910, 326911, 326914, 326915, 326916, 326918, 326919, 326920, 326921, 326922, 326923, 326924, 326926, 326928, 326931, 326932, 326935, 326937, 326940, 326947, 328228, 328230, 328231, 328232, 328233, 328245, 328246, 328248, 354732, 354733, 354734, 354735, 354736, 350484, 350487, 350488, 350489, 350490, 350491, 350492, 350493, 350494, 350196, 350497, 350498, 350512, 354710, 354711, 354712, 354713, 354714, 354715, 3564716, 354717, 354718, 354719, 354721, 354722, 354723, 354724, 354727, 354729, 354699, 354701, 354702, 354703, 354704, 354705, 354706, 354707, 354708, 354709, 328237, 328238, 328239, 328240, 328243, 329151, 329153, 329154, 329155, 354738, 369467, 369468, 369470, 369471, 369472, 369473, 369474, 369476, 369486, 369489, 369490, 369491, 369492, 369493, 369505, 369506, 369507, 322693, 322659, 322660, 322661, 322662, 322663, 322664, 322665, 322666, 322667, 322668, 322694, 322695, 282504, 282505, 282506, 282507, 282508, 282509, 282512, 282513, 283230, 283231, 321872, 321873, 321874, 401157, 401158, 401159, 401160, 401161, 401162, 401163, 401165, 401166, 401167, 401168, 401169, 401171, 401172, 401173, 401174, 401175, 401176, 401177, 401178, 401179, 401180, 401181.
Field Museum of Natural History (Chicago, USA)
103308, 103311, 103312, 103313, 103314, 103315, 103317, 103318, 103319, 103321, 103322, 103323, 97234, 97238, 97227, 97231, 97232, 97249, 97214, 97216, 97220, 97221, 97223, 97248, 111954, 111655, 153162, 153161, 179005, 179008, 87612, 87618.
British Museum of Natural History (London, UK)
19.7.7.2356, 19.7.7.2808, 47.508, 47.514, 47.510, 47.518, 47.509, 47.513, 95.350, 47.523, 47.529, 66.2584, 471537, 471535, 471538, 1979.2052, 1968.949, 1968.947, 1968948, 206814, 1957.10, 206813, 846146, 1023243, 51311, 196124, 196125, 54247, 23241.
The Musée National d’Histoire Naturelle (Paris, France)
1997-403, 1957.1340, 1955.21, 1957.519, 1953.4230, 1957.1036, 1957.1039, 1957.1043, 1953.425, 1958.104, 1957.1042, 1957.1037, 1957.1038, 1991.1220, 1952.532, 1957-329, 1957-325, 1957-314, 1958-63, 1957-321, 1957-528, 1957-316, 1958-61, 1958-62, 1957-825, 1957-318, 1957-328, 1957-330, 1966-1054, 1957-823, 1957-311, 1957-315, 1957-309, 1957-307, 1957-312, 1957-320, 1958-6423, 1957-306, 1957-326, 1957-323, 1958-65, 1957-322, 1958-107, 1958-106, 1957-1354, 1995-1700.
List of type specimens included in this study.
| Taxon | Synonym | Type locality | Collection |
|---|---|---|---|
| M. c. perpallidus | M. c | Egypt, Wadi Gedeed, Cairo, 4 km W; Cairo-Alexandria rd | FMNH |
| M. c. asyutensis | M. c | Egypt, Asyut, 13 mil SE; Wadi el Asyuti | FMNH |
| M. c. pallidus | M. c | Sudan, Atbara | BMNH |
| M. c. tripolidus | M. c | Libya, Gebel Limhersuk | BMNH |
| M. c. charon | M. c | Iran, Khuzestan, Ahwaz | BMNH |
| M. c. ismahelis | M. c | Arabia, Hufuf | BMNH |
| M. c. pelerinus | M. c | Arabia, Tebuk | BMNH |
| M. c. tuareg | M. c | Niger, Teguidda, near Tisem, west of Asben | BMNH |
| M. c. longifrons | M. c | Arabia, Jedda | BMNH |
| M. caudatus amplus | M. l | Libya, Fezzan, Gatrun | USNM |
| M. caudatus luridus | M. l | Libya, Cyrenaica, Bahr El Tubat, 21 KmE, SE Giarabub | USNM |
| M. erythrourus farsi | M. l | Iran, Fars, Bariz, 3 Km N, 50 Km N of Lar | USNM |
| M. l. aquilo | M. l | Zungaria, Gutschen | BMNH |
| M. l. gaetulus | M. l | Algeria | BMNH |
| M. l. caudatus | M. l | Libya, Tripolitania, Bir Ferdjan | BMNH |
| M. l. schouesboeii | M. l | Algeria, Tieret, Tibremt | BMNH |
| M. s. edithae | M. l | Arabia, Hufuf, Khudud Spring | BMNH |
| M. s. evelynae | M. l | Arabia, Hufuf, Khorassan Spring | BMNH |
| M. syrius | M. l | Syria, Al Qariatayn, Syrian Desert | BMNH |
Synonyms are according to Musser and Carleton (2005). Species codes are as follows: M. c=Meriones crassus and M. l=M. libycus. Collection codes are as follows: USNM, the Smithsonian Natural Museum of Natural History; FMNH, the Field Museum of Natural History; BMNH, the British Museum of Natural History.
Overview of sampling localities of Meriones crassus (M. c.) and M. libycus (M. l) used in this study.
| Taxon | Country | Locality | Abbreviation | Latitude | Longitude | Sample size | ||||
|---|---|---|---|---|---|---|---|---|---|---|
| F | M | Total | ||||||||
| M. c | Iran | Abas abad | ABA | 36.38 | N | 53.05 | E | 7 | 9 | 16 |
| M. c | Iran | Esfahan | ESF | 33.22 | N | 51.68 | E | 2 | 1 | 3 |
| M. c | Iran | Kashan | ESK | 33.98 | N | 51.45 | E | 0 | 2 | 2 |
| M. c | Iran | Kerman | KER | 30.28 | N | 57.10 | E | 8 | 9 | 17 |
| M. c | Iran | Khash, Iranshahr | KHS | 28.22 | N | 61.20 | E | 2 | 2 | 4 |
| M. c | Iran | Kashmar | KSH | 35.17 | N | 57.42 | E | 5 | 11 | 16 |
| M. c | Iran | Dasht-e- Lut | LUT | 32.58 | N | 59.03 | E | 1 | 1 | 2 |
| M. c | Iran | Mashad | MSH | 36.05 | N | 58.85 | E | 4 | 1 | 5 |
| M. c | Iran | Qazvin | QZV | 36.81 | N | 51.05 | E | 1 | 1 | 2 |
| M. c | Iran | Sabzevar | SBZ | 36.25 | N | 57.75 | E | 2 | 3 | 5 |
| M. c | Iran | Shahrud | SHD | 36.42 | N | 54.96 | E | 4 | 6 | 10 |
| M. c | Iran | Torbat-e-heidariyeh | TOH | 35.36 | N | 59.22 | E | 9 | 7 | 16 |
| M. c | Iran | Zahedan | ZHD | 29.21 | N | 60.87 | E | 2 | 5 | 7 |
| M. c | Iran | Mahallat | MHL | 33.75 | N | 50.50 | E | 1 | 1 | 2 |
| M. c | Iran | Tazuki | ZHD | 30.40 | N | 61.14 | E | 1 | 0 | 1 |
| M. c | Iran | Andimeshk | AND | 33.20 | N | 48.25 | E | 3 | 11 | 14 |
| M. c | Iran | Ahvaz | AVZ | 31.18 | N | 49.60 | E | 1 | 2 | 3 |
| M. c | Iran | Qhasr-e-shirin | KQA | 34.51 | N | 45.58 | E | 4 | 4 | 8 |
| M. c | Iran | Taj maleki | TJM | 29.76 | N | 50.56 | E | 2 | 0 | 2 |
| M. c | Afghanistan | Qala Bist | BST | 31.33 | N | 64.20 | E | 3 | 9 | 12 |
| M. c | Pakistan | Baluchistan | BAL | 28.58 | N | 65.41 | E | 1 | 4 | 5 |
| M. c | Pakistan | Nok kundi | NKK | 28.81 | N | 62.77 | E | 5 | 3 | 8 |
| M. c | Pakistan | Panjgur | PNG | 26.75 | N | 64.00 | E | 1 | 3 | 4 |
| M. c | Iraq | Kirkuk (Mulla Abd Allah) | IRQ1 | 35.47 | N | 44.13 | E | 5 | 3 | 8 |
| M. c | Iraq | Ali Garbi (Ali al Gharbi) | IRQ2 | 32.46 | N | 46.68 | E | 6 | 7 | 13 |
| M. c | Kuwait | Al ahmadi | KWT | 29.07 | N | 48.08 | E | 4 | 6 | 13 |
| M. c | Saudi Arabia | Qariya, Hafar-al-Batin | ARB | 27.20 | N | 44.00 | E | 4 | 6 | 10 |
| M. c | Saudi Arabia | Jeddah | JED | 21.51 | N | 39.22 | E | 0 | 1 | 2 |
| M. c | Israel | Ein Hussub | IRS | 31.78 | N | 35.22 | E | 1 | 2 | 3 |
| M. c | Jordan | Zarqa, Ma’am | JOR | 31.59 | N | 37.24 | E | 2 | 0 | 2 |
| M. c | Egypt | Al Qahriah | EGY | 30.10 | N | 31.33 | E | 8 | 4 | 12 |
| M. c | Libya | Awbari, Fezzan, El Gatrun | LIB | 29.06 | N | 15.78 | E | 8 | 5 | 13 |
| M. c | Mauritania | Fort Gouraud | MAU | 22.61 | N | -12.73 | W | 15 | 8 | 23 |
| M. c | Western Sahara | Aguera | MAU | 20.83 | N | -17.09 | W | 1 | 0 | 1 |
| M. l | Iran | Geno | GNO | 27.41 | N | 56.18 | E | 0 | 0 | 1 |
| M. l | Iran | Jajarm | JJM | 36.93 | N | 56.36 | E | 0 | 0 | 1 |
| M. l | Iran | Kashmar | KSH | 35.18 | N | 57.41 | E | 0 | 0 | 1 |
| M. l | Iran | Sabzevar | SBZ | 29.60 | N | 52.51 | E | 0 | 0 | 1 |
| M. l | Iran | Bajestan | BJS | 34.51 | N | 58.18 | E | 0 | 1 | 2 |
| M. l | Iran | Kouhak, Zabol | KOH | 31.06 | N | 61.75 | E | 2 | 0 | 2 |
| M. l | Iran | Garmsar | GAR | 35.13 | N | 52.18 | E | 0 | 0 | 3 |
| M. l | Iran | Kerman | KER | 29.90 | N | 56.53 | E | 0 | 0 | 3 |
| M. l | Iran | Gonbad-e-Kavus | GON | 37.23 | N | 55.08 | E | 2 | 3 | 6 |
| M. l | Iran | Shirvan | SHV | 37.45 | N | 57.91 | E | 0 | 3 | 4 |
| M. l | Iran | Torbat-e-heidariyeh | TOH | 35.28 | N | 59.21 | E | 0 | 1 | 4 |
| M. l | Iran | Zahedan | ZHD | 29.53 | N | 60.83 | E | 3 | 2 | 5 |
| M. l | Iran | Torbat-e-Jam | TOJ | 35.21 | N | 60.61 | E | 0 | 1 | 8 |
| M. l | Iran | Nehbandan | NEH | 31.53 | N | 60.03 | E | 1 | 4 | 8 |
| M. l | Iran | Ghoochan | SHV | 37.10 | N | 58.50 | E | 3 | 5 | 8 |
| M. l | Iran | Sabzevar | SBZ | 36.20 | N | 57.71 | E | 3 | 4 | 8 |
| M. l | Iran | Khash | KHS | 35.18 | N | 58.45 | E | 2 | 6 | 9 |
| M. l | Iran | Mashhad | MSH | 36.26 | N | 59.61 | E | 1 | 3 | 10 |
| M. l | Iran | Sarakhs | TND | 36.50 | N | 61.05 | E | 0 | 5 | 9 |
F, female; M, male; Total, both sexes including unknown sex specimens.
Definition and numbering of the landmarks describing the ventral skull morph.
| Landmark | Definition |
|---|---|
| 1 | Rostral tip of internasal suture |
| 2 | Most lateral junction point of incisive alveolus and body of premaxillary bone |
| 3 | Most rostral point of foramen magnum |
| 4 | Most lateral point of occipital condyle |
| 5 | Most caudal point of acoustic tympanic bulla |
| 6 | Rostral curvature point at level of the meatus |
| 7 | Most lateral point of zygomatic arch at maximum width of skull |
| 8 | Intersection of zygomatic arch and vertical line passing through most caudal point of third molar |
| 9 | Rostral point of zygomatic plate |
| 10 | Maximum curvature of zygomatic plate in infraorbital foramen |
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