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THE QUARTERLY REVIEW of BIOLOGY VOL. 43, NO. I March, 1968 THE QUARTERLY REVIEW of BIOLOGY LIFE CYCLE ORIGINS, SPECIATION, AND RELATED PHENOMENA IN CRICKETS BY RICHARD D. ALEXANDER Museum of Zoology and Departmentof Zoology The Universityof Michigan,Ann Arbor ABSTRACT Seven general kinds of life cycles are known among crickets; they differ chieff,y in overwintering (diapause) stage and number of generations per season, or diapauses per generation. Some species with broad north-south ranges vary in these respects, spanning wholly or in part certain of the gaps between cycles and suggesting how some of the differences originated. Species with a particular cycle have predictable responses to photoperiod and temperature regimes that affect behavior, development time, wing length, bod)• size, and other characteristics. Some polymorphic tendencies also correlate with habitat permanence, and some are influenced by population density. Genera and subfamilies with several kinds of life cycles usually have proportionately more species in temperate regions than those with but one or two cycles, although numbers of species in all widely distributed groups diminish toward the higher lati­ tudes. The tendency of various field cricket species to become double-cycled at certain latitudes appears to have resulted in speciation without geographic isolation in at least one case. Intermediate steps in this allochronic speciation process are illustrated by North American and Japanese species; the possibility that this process has also occurred in other kinds of temperate insects is discussed. INTRODUCTION the Gryllidae at least to the Jurassic Period (Zeuner, 1939), and many of the larger sub­ RICKETS are insects of the Family families and genera have spread across two Gryllidae in the Order Orthoptera, or more continents. and are identifiable by their jump­ Crickets occupy today all of the major ing hind legs, three tarsal segments, terrestrial habitats between approximately lati­ C and long tactual cerci bearing tudes 55° N and 55° S, or, roughly, the region clumps of knobbed hairs. They range in body between the boundaries of the northern and length from l or 2 mm to over 50 mm. The southern coniferous forests, or taiga (Fig. l). family comprises some 2455 known species, In the course of achieving this broad distribu­ presently arranged in 14 subfamilies and about tion, they have evolved at least seven kinds of 333 genera (Table I). Fossil evidence dates life cycles, most of them more than once in 1 2 THE QUARTERLY REVIEW OF BIOLOGY [VOLUME 43 various parts of the world. These different .!,"Cl .C:"t:1 ........ -i:: "'01 bO adjustments to climatic variation involve far 0 01 -~; fl d d "'-bl) "[t more than the simple synchronizing of resis­ ...~ ::Id l~ V -a-01 g ..o.~ ~"'..0 8 tant life stages with harsh seasons. Within vO 01 .C: d v~ <I.I 5l each species studied, there are also genetically u"' 01 s- ._,· C ~ ti ..0Ei "'V .,0 !'.l01 V and environmentally induced variations in 01 C: ..,"',2 0 8 0 t .., .... ."Cl j:l morphology, physiology, and behavior that are ::;; ~ ...... 0 ti si "Cl bO consistent with life cycle pattern, latitude, and < .....~ d "' ..: '\s ~ 0.. < -~ ., ..,-~i-s ..9 habitat. Particularly during the past two dec­ :c i·; ";' ~ ades, the nature of many of these phenomena, u .s~:a., ii.,o bl) ~ -~ '\? ;,.,.-~ ..0 § "o ... ~ including some of their probable origins and s . ~,d M """"o :::,"t:I 01 <I.I ., histories, has been clarified by parallel and ;a-~ ~ f;!·> v· v"t:I ~ v"t:I d O 01 e!'S; complementary investigations in Europe, 01 0 - ~~ s..8 ~s] - ., bl) s ...... Japan, Australia, and North America. (J This paper summarizes the available infor­ gd ,..;"0-i mation and compares variously related species z -~o..-~ 01 se .., 9 .8 ""; d.., in an attempt to reconstruct some of the evo­ --~ 'ii ;:;s 0.. .c: l::., -~ 8 ~ .., 0:, ... ·;:: <I.I <I.I 8 fa<:::, oz lutionary changes by which known combina­ z~ "t;I ~ :g :::, • =· tions of characteristics have come about. ~~ ~..:I .;:f': d 01 El -~ o..- -~= ~ gjg " -6 ~ ... d g-is :g ~ ~ ] 0 O;..: :E~ 8·t ... ::I KINDS OF LIFE CYCLES ~... ~ ~~ zo ..:1..: \..'.) Crickets are oviparous, hemimetabolous in­ -s" ,e .,; sects. Their eggs are injected into soil, plant ._ .., ..0..,... C .c: ..,"' .c: 01 01 stems, or twigs, or laid in burrows. There are fa< -j:l -£ ...l ;:;~ !'.i from five to twelve juvenile instars, or molt ~ -~ .., t:: .c:., .c:!'.i ,; < ..,z ~< ::I ,,, ... ~~ u stages, and the juveniles resemble the adults, r"' l ... ::I <I.I ~ ~ " :c ..0 ..0 ~ -eG ... live in the same general habitats, and eat the .£:) ,;; fl):.= ~~ ~t ..,;:! ..,u <I.I- _v same kinds of food. Most species studied seem l:: .........~ ~ ·s.E..,...... ]~ to eat either plant or animal food of a wide -s" ,,, variety; some tree crickets are believed to sub­ C ,,; el 0 0 - 0 <J:) sist largely on aphids, and some mole crickets .~ u 8 $ ,, z <J:) .;:,.. ; ~ on roots and earthworms. C .: ~ The seven general kinds of cricket life ~ ~ x cycles, six of which are illustrated in Figures 2 ~ ~ ~ .., C<") 0 ...... C<") and 3, are the following: z ~ .... <J:) "'< .., (l) Continuous breeding without any diapause "' c.; in environments without marked seasonal changes. Examples are Acheta domesticus (Lin­ .., !!l naeus), Gryllodes sigillatus (Walker) (species ~ !!l < <I.I ~ that are found around the world near human z ~u .::1... .., ,,, ·.:: u habitations, but that otherwise occur only in !!l u "t;I <I.I ti "t;I s ~ tropical regions); Gryllus assimilis Fabricius in O' ..:.: u <I.I d i -~ ·.:: OJ the West Indies, Mexico, and extreme southern ..,s u u ·a. ...l 0 er, <I.I ., bO United States (Florida, Texas, California); u <I.I -5i d ... ::I 0 Teleogryllus mitratus (Burmeister) in tropical !-< ~ ~ ...l Asia (Masaki and Ohmachi, 1967); and Tohila atelomma Hubbell, a cave inhabitant in Yu­ OJ >, <I.I 01 catan (Hubbell, 1938). [When references are .., .., <I.I 01 01 d .s est e ·;::: ,,, not given, life cycle determination is based on i i:: ·;::: Cl.. < i: 0 my studies. I have reared all species mentioned "' :g <I.I .., bl) ::,= i:: 0.. u d except Tohila atelomma, Myrmecophila ne­ 01 5 "' u 8 d -; <I.I d .., .c: brascensis, Gryllotalpa gryllotalpa, and those ..0 ..fa< ..i:i.. i:i.. :a~ n .....= TABLE I-Continued ! Scleropterinae Stiff-Winged Crickets 3 16 soil surface, leaf Palearctic, Ethiopian, medium to large; wings hard, horny, litter Oriental, Australian and opaque r, ~ Pteroplistinae Feather-Winged Crickets I 3 low herbs and water Oriental, Australian tiny, soft-bodied t'l surface C) "Trigonidiinae Sword-Tailed Crickets 28 380 trees, bushes, herbs, World-wide small or tiny, delicate, usually pale r,~ leaf litter brown in color t'l .. 0 Cachoplistinae None I 4 soil surface? Ethiopian, Oriental, tiny, soft-bodied :,;;, Australian ..... "Mogoplistinae Short-Winged Crickets 14 200 bushes, herbs, ground World-wide small or tiny, short forewings, en- litter larged pronotum, body with scales ~ ::i.. "Myrmecophilinae Ant-Loving Crickets 5 45 ant nests World-wide tiny, wingless, hump-backed, no audi- § tory organs, short antennae, reduced eyes. C)~ "Nemobiinae Ground Crickets 16 200 soil surface World-wide small, cf with tibial or tegminal gland s;: "Gryllinae Field and House Crickets 70 550 soil surface and World-wide large size, yellow to black, ~ with ::j shallow burrows long ovipositor 0 ~ "Brachytrupinae Short-Tailed Crickets 12 42 soil surface and deep World-wide resemble Gryllinae except with short burrows (except Europe) or no external ovipositor ~ Q •Gryllotalpinae Mole Crickets 6 65 subterranean World-wide mostly large, heavy bodied; shovelling ..... forelegs, pointed head, cylindrical C) body, dense body setae ~ Totals 14 333 2455 ~ • Subfamilies discussed in this paper. <:.>o 4 THE QUARTERLY RETIIEW OF BIOLOGY [VOLUME 43 0 0 !:e !Q 0 ~ 0 (X) 0 0 <:t 0 (X) 0 ~ 0 !£ 0 (t) from Asia and Africa, and I have studied vir­ sis Luggar, in Austin, Texas: "During April tually all North American species.] This kind and May the testes, in active spermatogene­ of cycle is rarer than might be supposed. Many sis... " and "At this same period of the tropical and sub-tropical crickets are evidently year the abdomen of the female Myrmecophila cyclic in association with dry and wet seasons. is found to contain a few very large elliptical Wheeler (1900) even implies seasonality in the white eggs. ." ant nest inhabitant, Myrmecophila nebrascen- 2. One generation each year with egg dia- MARCH 1968] LIFE CYCLE ORJGlNS AND SPECIATION IN CRICKETS 5 Jan Feb vernalis fu/toni veletis pennsylvanicus firmus rubens assimilis FIG. 2. DIAGRAMS OF THE LIFE CYCLES OF THE SEVEN FIELD CRICKETS (Gryllus spp.) IN EASTERN NORTH AMERICA, ILLUSTRATING FOUR OF THE SEVEN GENERAL KINDS OF LIFE CYCLES KNOWN IN CRICKETS Gryflotalpa grylIota/pa Nemobius sylvestris FIG. 3. DIAGRAMS OF THE Two-YEAR LIFE CYCLES REPORTED FOR Gryllotalpa gryllotalpa IN SPAIN (Morales, 1940) AND Nemobius sylvestris IN ENGLAND (Gabbutt, 1959) G THE QUARTERLY REVIEW OF BIOLOGY [VOLUME 43 pause during winters or dry seasons. Examples life cycle in tropical India for the univoltine are Gryllus pennsylvanicus (Burmeister), Mio­ burrowing cricket Brachytrupes achatinus Stoll, gryllus verticalis (Serville), Valerifictorus mi­ which may be related to this one. Most individ­ cado (Saussure) (introduced) in North America; uals mature in June, but eggs are not laid un­ all North American Nemobiinae north of til September or October; thus juvenile growth about the 30th parallel; most temperate, vege­ is retarded during winter, and the summer dry tation-inhabiting Gryllidae (involving three sub­ season is spent in a prolonged adult stage. families: Oecanthinae, Trigonidiinae, and Ene­ (6) Two generations each year with juvenile opterinae); Teleogryllus commodus (Walker) diapause during winters. Examples are Gryl­ in Australia (Hogan, 1960), T. emma (Ohmachi lus rubens Scudder and G.
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