(MAMMALIA: RODENTIA) Igor Ya. Pavlinov
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Igor Ya. Pavlinov A REVIEW OF PHYLOGENY AND CLASSIFICATION OF GERBILLINAE (MAMMALIA: RODENTIA) ЗООЛОГИЧЕСКИЕ ИССЛЕДОВАНИЯ № 9 Зоологический музей МГУ Zoological Museum of Moscow State University 1 Çîîëîãè÷åñêèå èññëåäîâàíèÿ N 9 Ïîñâÿùàåòñÿ ïàìÿòè Â. Ã. Ãåïòíåðà È. ß. Ïàâëèíîâ ÎÁÇÎÐ ÔÈËÎÃÅÍÈÈ È ÊËÀÑÑÈÔÈÊÀÖÈÈ GERBILLINAE (MAMMALIA: RODENTIA) ÈÇÄÀÒÅËÜÑÒÂÎ ÌÃÓ ÌÎÑÊÂÀ – 2008 2 Zoologicheskie Issledovania N 9 To the memory of W. G. Heptner Igor Ya. Pavlinov A REVIEW OF PHYLOGENY AND CLASSIFICATION OF GERBILLINAE (MAMMALIA: RODENTIA) MOSCOW UNIVERSITY PUBLISHING MOSCOW – 2008 3 ÇÎÎËÎÃÈ×ÅÑÊÈÅ ÈÑÑËÅÄÎÂÀÍÈß ¹ 9 ZOOLOGICHESKIE ISSLEDOVANIA ¹ 9 Editor-in-Chief: O. L. Rossolimo Editor of this issue: A. V. Sysoev Igor Ya. Pavlinov. A review of phylogeny and classification of Gerbillinae (Mammalia: Rodentia). — Moscow: Moscow Univ. Publ. 2008. 68 p. Under consideration are modern concepts of phylogeny and taxonomy of a rodent subfamily Gerbillinae (gerbils and jirds). Principal pathways of morphological evolution of the masticatory apparatus (mainly cheek teeth) and the ossean middle ear (mainly mastoid part of the auditory bulla) in Gerbillinae are characterized. Fossil genera referred to Gerbillinae are briefly reviewed, the genera Epimerionaes and Pseudomeriones are shown to be non-gerbillines. Predominating hypotheses about both phylogenetic position of Gerbillinae within Muroidea and genealogical relationships among its genera are outlined. Principal points of both agreements and disagreements between morphology-based and DNA-based phylogenies are indicated. Previous and current classifications of Gerbillinae are exposed, with most attention being paid to the one developed by the author. Suprageneric taxa and genera recognized in the latter are described: each taxon is characterized in respect to its major morphological features, principal fossils, various versions of its contents and relationships, and nomenclature upon a need. The identification keys for the genera and subgenera are provided. Ill. 26. Bibl. 133. Ãëàâíûé ðåäàêòîð: Î. Ë. Ðîññîëèìî Ðåäàêòîð âûïóñêà: À. Â. Ñûñîåâ È. ß. Ïàâëèíîâ. Îáçîð ôèëîãåíèè è êëàññèôèêàöèè Gerbillinae (Mammalia: Rodentia). — Ìîñêâà: Èçä-âî Ìîñê. óí-òà. 2008. 68 ñ. Ðàññìîòðåíû ñîâðåìåííûå ïðåäñòàâëåíèÿ î ôèëîãåíèè è ñèñòåìàòèêå ãðûçóíîâ ïîäñåìåéñòâà Gerbillinae (ïåñ÷àíêè). Îõàðàêòåðèçîâàíû îñíîâíûå íàïðàâëåíèÿ ñïåöèàëèçàöèè æåâàòåëüíîãî àïïàðàòà (ãëàâíûì îáðàçîì ù¸÷íûõ çóáîâ) è êîñòíîãî ñðåäíåãî óõà (ãëàâíûì îáðàçîì ìàñòîèäíîãî îòäåëà ñëóõîâîãî áàðàáàíà) ïåñ÷àíîê. Ïðåäñòàâëåí êðàòêèé îáçîð èñêîïàåìûõ ðîäîâ, îòíîñèìûõ ê Gerbillinae; ïîêàçàíî, ÷òî ðîäû Epimerionaes è Pseudomeriones äîëæíû áûòü èñêëþ÷åíû èç ýòîãî ïîäñåìåéñòâà. Èçëîæåíû îñíîâíûå ãèïîòåçû î ïîëîæåíèè ïîäñåìåéñòâà ñðåäè Muroidea, ñòðóêòóðà ôèëîãåíåòè÷åñêèõ ñâÿçåé ìåæäó åãî ðîäàìè; îòìå÷åíû îñíîâíûå òî÷êè ñîãëàñèÿ è ðàñõîæäåíèÿ ìîðôîëîãè÷åñêèõ è ìîëåêóëÿðíî- ãåíåòè÷åñêèõ ãåíåàëîãè÷åñêèõ ñõåì. Êðàòêî ðàññìîòðåíû ïðåæíèå è ñóùåñòâóþùèå êëàññèôèêàöèè Gerbillinae; îñíîâíîå âíèìàíèå óäåëåíî ñèñòåìå, ðàçðàáîòàííîé àâòîðîì. Îõàðàêòåðèçîâàíû ïðèçíàííûå â ýòîé ñèñòåìå íàäðîäîâûå ãðóïïû è ðîäû ñîâðåìåííûõ ïåñ÷àíîê: äëÿ êàæäîãî òàêñîíà óêàçàíû îñíîâíûå ìîðôîëîãè÷åñêèå ïðèçíàêè, êëþ÷åâûå èñêîïàåìûå ïðåäñòàâèòåëè, èçëîæåíû ðàçíûå âåðñèè ñîñòàâà è ðîäñòâåííûõ ñâÿçåé, ïðè íåîáõîäèìîñòè çàòðîíóòû âîïðîñû íîìåíêëàòóðû. Ïðåäñòàâëåíû êëþ÷è äëÿ îïðåäåëåíèÿ ðîäîâ è ïîäðîäîâ ñîâðåìåííûõ ïåñ÷àíîê. Èëë. 26. Áèáë. 133. ISSN 1025-532Õ © I.Ya.Pavlinov, 2008 © Zoological Museum, 2008 4 CONTENTS Introduction......................................................................................................7 Evolutionary trends and morphological features.............................................9 Bony elements of masticatory apparatus...............................................9 Dentition.............................................................................................14 Middle ear...........................................................................................16 Mastoideum..........................................................................16 Transmitting system.................................................................19 A review of fossil genera..............................................................................21 Modern phylogenetic hypotheses..................................................................25 Relationships to the other Muroidea....................................................25 Phylogenetic trees for the the Recent Gerbillinae...............................28 Taxonomy of Recent Gerbillinae...................................................................30 A review of principal classifications...................................................30 Tribe Taterillini....................................................................................33 Subtribe Taterillina s.str............................................................34 Subtribe Gerbillurina.................................................................38 Tribe Ammodillini...............................................................................40 Tribe Gerbillini s.str. ..........................................................................42 Subtribe Gerbillina s.str. ...........................................................43 Subtribe Desmodilliscina..........................................................47 Subtribe Pachyuromyina...........................................................48 Subtribe Rhombomyina............................................................50 Keys to the genera and subgenera.................................................................56 Key to the Asian taxa...........................................................................56 Key to the Northern and Eastern African taxa...................................57 Key to the Southern African taxa........................................................58 Conclusions....................................................................................................59 Acknowledgments..........................................................................................60 References......................................................................................................61 Summary........................................................................................................66 Index...............................................................................................................68 5 ÑÎÄÅÐÆÀÍÈÅ Ââåäåíèå.................................................................................................................................7 Ýâîëþöèîííûå òåíäåíöèè è äèàãíîñòè÷åñêèå ïðèçíàêè.................................9 Êîñòíûå ýëåìåíòû æåâàòåëüíîãî àïïàðàòà..................................................9 Çóáíàÿ ñèñòåìà.....................................................................................................11 Ñðåäíåå óõî...........................................................................................................16 Ìàñòîèä......................................................................................................16 Ïðîâîäÿùàÿ ñèñòåìà..............................................................................19 Îáçîð èñêîïàåìûõ ðîäîâ.........................................................................................21 Ñîâðåìåííûå ôèëîãåíåòè÷åñêèå ãèïîòåçû..........................................................25 Ñâÿçè ñ äðóãèìè Muroidea...............................................................................25 Ôèëîãåíåòè÷åñêèå ñõåìû äëÿ ñîâðåìåííûõ Gerbillinae.......................28 Ñèñòåìàòèêà ñîâðåìåííûõ Gerbillinae....................................................................30 Îáçîð îñíîâíûõ êëàññèôèêàöèé................................................................30 Òðèáà Taterillini......................................................................................................33 Ïîäòðèáà Taterillina s.str. ......................................................................34 Ïîäòðèáà Gerbillurina.............................................................................38 Òðèáà Ammodillini..............................................................................................40 Òðèáà Gerbillini.....................................................................................................42 Ïîäòðèáà Gerbillina s.str. .....................................................................43 Ïîäòðèáà Desmodilliscina....................................................................47 Ïîäòðèáà Pachyuromyina.....................................................................48 Ïîäòðèáà Rhombomyina......................................................................50 Êëþ÷è äëÿ îïðåäåëåíèÿ ðîäîâ è ïîäðîäîâ.........................................................56 Êëþ÷ äëÿ òàêñîíîâ Àçèè.................................................................................56 Êëþ÷ äëÿ òàêñîíîâ Ñ. è Â. Àôðèêè...........................................................57 Êëþ÷ äëÿ òàêñîíîâ Þ. Àôðèêè..................................................................58 Çàêëþ÷åíèå........................................................................................................................59 Áëàãîäàðíîñòè..................................................................................................................60 Ëèòåðàòóðà..........................................................................................................................61