Gerhard Storch – a Legend in Mammal Research
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Gerhard Storch – a legend in mammal research During the second half of the 20th century, mammalogy underwent a revolutionary burst in knowledge about diverse aspects of mammalian biology, phylogeny of particular clades and detailed information on the course of their history. It is ascribed primarily to extensive instrumentation innovations and to methods not available earlier. In the time of big data and excessive information boom, we somehow forgot that, in general, scientific progress results first of all from initial setup of new research fields, identification of links between different research areas and establishing conceptual outlines for research efforts to come. Without the intellectual efforts of personalities disposed to respond to these demands, no progress in any branch of science would be possible. The late Gerhard Storch, to whose memory the present volume is devoted, personified this in an exemplary way. He has been rightly denoted as one of the greatest mammalogists of our time. He was an outstanding integrative personality, who contributed to the shift in knowledge of both recent and fossil mammals in an essential way. Gerhard Storch giving a talk at the Congressus Theriologicus Internationalis in Brno, 1978 (photo by J. Červený) Gerhard Storch was born on May 21st, 1939 in Frankfurt am Main, where he also grew up and spent most of his life. As an enthusiastic young naturalist, he became already at the age of 15 a member of the Senckenberg Gesellschaft für Naturforschung. There began his close co-operation with Heinz Felten (1922 – 2000), then a research fellow of the Senckenberg Resarch Institute and Natural History Museum Frankfurt, and experienced expert in small ground mammals and bats. Gerhard started with field study of these groups, in order to resolve diverse questions of their biology, and, first of all, the taxonomic, biogeographic and evolutionary settings of particular clades. During the 1960s, he undertook about 50 field trips to study native rodents and insectivores in the wider Rhine-Main area, and particularly in the Alps, as the major crossroad of the mid-European mammalian biodiversity. Detailed investigations of small Alpine mammals were resumed in 1971, with collective trips to the Italian Alpine region, resolving the status of enigmatic endemic forms of that region [44, 112, 189]. Simultaneously, since the beginning of 1960s, he regularly conducted research trips into exotic destinations of the Mediterranean and Middle East, regions which mammal fauna was then nearly unknown. These adventures were undertaken first with Franz Malec, a friend of his age and with similar interests (later a director of the Natural History Museum in Kassel), and later in the sixties, as part of a common large-scale research program, also with Heinz Felten, Friedericke Spitzenberger and Dieter Kock. The systematic field study of the mammal fauna in the Balkans, Iran, Turkey, Syria, Spain, Italy, Mediterranean v islands and northern Africa resulted in a series of papers, which present true pillars of knowledge on mammal fauna and biogeography of these regions [1–5, 9, 10, 16–21, 23–25, 28, 39–44, 62, 68, 234]. They resolved numerous uncertainties about the taxonomic and distribution status of particular local forms (with descriptions of several new taxa [1, 9, 16, 42]), and at the same time established a high standard for further investigations of that kind. The refined techniques of comparative analyses combining detailed morphometrics, scoring of non-metric variables and contextual factors (habitat requirements etc.) are shown here as tools for disclosing the patterns of interspecific and intraspecific variations. This also enabled to address the questions of biogeographic and historical factors influencing local faunal development. It is quite illustrative, in this respect, that beginning with their first expeditions, Storch and Malec combined the large-scale collecting of extant mammals with increasing interest in the study of subfossil and fossil mammals, and practical exploration of suspected fossiliferous sites. In his search for the historical roots of extant taxa and the dynamics of extant communities, Gerhard focused his attention first on the development of the Late Pleistocene and Holocene fauna and excavations of cave deposits of that age, including various archaeological sites both in Germany [14, 29, 35, 36, 56–58, 63–64, 72, 76, 88, 119, 130, 152, 224] and abroad [15,16– 18, 23–26, 31–32, 37, 41, 62, 99]. This brought him to compile the first comprehensive summaries on the history of mammal communities along the Pleistocene / Holocene transition in Central Europe, Northeast Iran and South Anatolia [35, 62, 99], highlighting phenotype shifts in some taxa [14, 17, 36], and the geographical variations of these phenomena [130, 224]. The broad span of Gerhard’s interests at the time of his university studies (in Darmstadt, Vienna and Frankfurt) was further expanded after he managed to get a temporary position as research assistant at the Senckenberg Mammalogy Section in 1967 (funded by the Deutsche Forschungsgemeinschaft – DFG). At that time, he focused his work on a detailed comparative study of the functional and morphological aspects of mastication in diverse mammalian clades [8, 11, 13]. The monographic survey of bats [11], an essential part of his PhD thesis (1967), is particularly worthy of attention. It integrated an enormous amount of myological and craniomorphological data, biomechanical models and recordings of jaw kinematics, with details of feeding biology and hypotheses on phylogenetic relations among individual clades of the order. The study provided a complex view on pathways of feeding specializations, and among other things, explicitly demonstrated the monophyly of the order, long before molecular tools resolved doubts on that point. In 1969, Gerhard Storch became a fulltime researcher at the Senckenberg Research Institute and Natural History Museum Frankfurt, and thanks to the initiative of Heinz Felten, the head of the newly established Section Mammalogy II, a research unit focused on fossil mammals. The topic of his research was Tertiary and Quaternary small mammals (rodents, insectivores, bats). Besides continuous interest in the Holocene and Late Pleistocene interface of neontologic topics, and the palaeobiogeography of Mediterranean mammals, Gerhard extended the frame of his investigations to mammal faunas of the Early Pleistocene, Pliocene and Late Miocene age. During the seventies, in co-operation with Franz Malec and Jens Franzen, he described rich assemblages from Hohensülzen [30] and Dorn-Dürkheim [38, 40, 46, 55, 157, 199], as well as Early and Middle Pleistocene sites on the islands of Malta, Chios and Kalymnos [34, 39, 45]. By the end of the 1970s, he took care of a unique collection of mammalian fossils from one of the world’s most important Lagerstätte: the early middle Eocene Messel Pit fossil site. In 1978, he described from there the earliest member of the order Pholidota: Eomanis waldi [47, 54, 65, 95, 135]. At the same time, he paid special attention to one of his favorite groups, particularly well-represented in the Messel fossil record – bats. Thanks to abundant and extraordinary well-preserved material from Messel, he subsequently described (partly together with J. Habersetzer, G. Richter, J. Smith or B. Sigé) several new clades of early bats (including a new family Hassianycteridae), revealed unexpected details on their diet, dental characters, diversification of their wing morphology, echolocation capacities, foraging strategies and pathways of the earliest adaptive radiations within the order [60–61, 84, 91, 105–106, 108, 113, 124–126, 131, 141, 151, 179, 182, 217]. No wonder that Gerhard Storch is widely remembered as one of the most prominent experts in that group and its history. Analysing further Messel fossils, he contributed in a similar way to the knowledge of the early history of a wide spectrum of mammalian clades. For instance, his description of Eurotamandua joresi [67, 96] started vivid discussions on the relationships between Afrotheria and Xenarthra, and extensive reconsideration of Eocene mammal radiations, prefiguring the current concept of Atlantogenata [121, 209, 211, 213, 218–220, 240]. An extraordinarily preserved specimen of Eogliravus wildi, the earliest dormice rodent, elucidated phylogenetic relationships of that family [198, 200, 216, 242]. His work on the insectivorous mammals of Messel, both placentals and marsupials [71, 78, 80, 85–87, 92–93, 98, 132, 133, 145–148, 177, 178] helped defining the diagnostic characters and understanding the early diversification of several clades (e.g. Proteutheria, Lipotyphla, Primates), and the palaeobiological factors of their radiations. Storch’s extraordinary capacity for synthesizing complex results enabled him to compile a series of studies in which the distinguishing features of the Messel fauna and its significance for solving palaeobiogeographic and palaeoenvironmental questions in the Eocene were demonstrated in detail [75, 81–83, 92–97, 100, 109–110, 116, 127, 131–137, 140, 142, 144, 158, 183, 210, 212, 233, 241, 245]. Likewise, Storch paid attention to other early Paleogene fossils, particularly from Geiseltal [78, 111, 140, 158, 208], the Paleocene site Walbeck [256], and the eomyid rodents from the Oligocene site Enspel, which revealed the earliest gliding adaptation in rodents [162, 173, 193]. While he worked with continuous efforts on solving enigmas about the Paleogene stage