Michael R. Maxwell’s research while affiliated with University of San Diego and other places

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Publications (13)


Fitness effects of nuptial gifts in the spider Pisaura mirabilis: Examination under an alternative feeding regime
  • Article
  • Full-text available

November 2018

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152 Reads

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9 Citations

Journal of Arachnology

Michael R. Maxwell

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Nuptial feeding has variable effects on fitness within a species, partly driven by variation in female diet. We investigate nuptial feeding in the spider Pisaura mirabilis (Clerck, 1757) under a feeding regime that has not been explored: Starvation after mating and gift consumption. We vary gift size and gift number to examine the effects on mating behavior and components of female fitness. With regard to gift size, copulation duration increased with larger gift size, but no component of female fitness was affected (time to oviposition, egg sac mass, female lifespan). These results corroborate other examinations of gift size in P. mirabilis. Given a likely male benefit (prolonged copulation) for larger gift size and no demonstrated female benefit, sexual conflict stands as a likely explanation for male benefits due to large nuptial gift size. With regard to gift number, components of female fitness were not affected by the consumption of one or two extra gifts. This agrees with other studies, although we note that some experiments have found the consumption of extra gifts to increase female fitness. As for males, they were more likely to copulate when they had gifts, as in other studies. We conclude some support for sexual congruence with regard to gift number, as males and females stand to benefit simultaneously from the mere presence of the gift, and females might benefit from the consumption of multiple gifts. Thus, both sexual conflict and sexual congruence appear to be at work regarding the evolution of nuptial gifts in Pisaura mirabilis.

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Female predatory response to conspecific males and heterospecific prey in the praying mantis Mantis religiosa: evidence for discrimination of conspecific males

February 2016

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331 Reads

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2 Citations

Journal of Ethology

Sexual cannibalism, the attack and consumption of the opposite sex by a conspecific during courtship, copulation, or soon after copulation, is widespread among invertebrates, but the causes of this behavior are not fully understood. We examine the mistaken identity hypothesis, which posits that non-copulatory cannibalism occurs because females do not recognize conspecific males as potential mates. This hypothesis predicts indiscriminate predatory behavior by females towards conspecific males and heterospecific prey. This prediction remains largely unexamined. We tested this prediction in the praying mantis Mantis religiosa (Mantodea: Mantidae) through two behavioral experiments. In experiment 1, we presented targets in isolation to females: dead conspecific males or dead common prey items from the field (grasshopper Chrysochraon dispar). In experiment 2, we presented simultaneous live targets to females: conspecific males and grasshoppers. In both experiments, we varied female mating status (virgin vs. mated). Results indicate some degree of attack discrimination by the females. In experiment 1, the females were significantly more likely to strike at the grasshoppers, and showed a shorter latency to strike at grasshoppers. In experiment 2, females tended to preferentially strike at the grasshoppers, although this result fell short of statistical significance. Female discrimination between the males and grasshoppers may be explained, in part, by the size difference between the males and grasshoppers. Female mating status did not affect the occurrence or latency to strike in either experiment. These results fail to strongly support the prediction of indiscriminate predatory behavior by females, while contributing to a broader taxonomic assessment of the mistaken identity hypothesis.


Developmental Patterns in Stagmomantis limbata (Mantodea: Mantidae): Variation in Instar Number, Growth, and Body Size

July 2014

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114 Reads

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12 Citations

Journal of Orthoptera Research

Development affects many components of life history and fitness, including body size. The present study examined the influence of developmental pattern, specifically the number of nymphal instars, on body size (pronotum length) in the praying mantid Stagmomantis limbata Hahn. Mantids were reared in the laboratory from hatching, on standardized diet, to examine variation in instar number. These lab data were then used to assess developmental patterns forfield-collected female nymphs. Laboratoryreared males and females varied in number of instars. Most females required 6 nymphal instars to reach adulthood (64%), whereas 36% underwent 7 instars. Seven-instar females reached the 4th, 5 th, and 6th instars faster than six-instar females, but had shorter pronota than the six-instar females at each of these stages. Seven-instar females were longer than six-instar females at adulthood. Interestingly, the total developmental period from hatching to adulthood was similar for lab-reared seven-instar and six-instar females. In the lab, most males (91%) underwent 6 instars, with the remaining 9% following a five-instar pattern. By the 4th instar, differences between the sexes began to appear. From the 4th instar onwards, females typically took less time than males to reach each instar. From the 5th instar onwards, females were longer than the males, and were longer as adults. Variation in developmental pattern (number of instars) was evident among siblings from the same ootheca; such intra-clutch variability in number of instars may be a bet-hedging strategy by ovipositing females in a variable environment. The laboratory data allowed for the detection of six-instar and seven-instar patterns among the field-collected females. The field-collected data suggest that females undergoing 6 nymphal instars reach adulthood later in the season, and at smaller body size, than seven-instar females.


A synoptic review of the genus Stagmomantis (Mantodea: Mantidae)

February 2014

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224 Reads

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21 Citations

Zootaxa

Stagmomantis is a remarkable genus within the Mantodea, being relatively species-rich and geographically widespread. Yet, the number of species within the genus remains curiously unresolved. The present synoptic review surveys the literature on Stagmomantis to identify named species for which scientific consensus exists, as well as to summarize basic biological information for each species, including geographic distribution, morphological features, and sex-specific biometric data. The review identifies 23 consensus taxa within Stagmomantis: 22 separate species, with one of these species, S. montana, split into two subspecies (S. m. montana and S. m. sinaloae). The review indicates morphological features that may prove to be diagnostic for a given species, particularly when examined in conjunction with male genitalia. Such features include dark spots on the anterior femur (S. amazonica, S. centralis, S. marginata, S. nahua, S. venusta, S. vicina), spines or denticulations on the anterior coxa (S. colorata, S. montana montana, S. parvidentata, S. theophila), and dark bands on abdominal tergites (S. californica, S. colorata, S. domingensis). Color variation of certain features with respect to body coloration, such as stigma coloration and body and leg markings, requires more attention. Information on life history, reproduction, and ecology are summarized, particularly for temperate populations of S. carolina and S. limbata. While the 23 consensus taxa represent a robust appraisal of the existing literature, some taxonomic uncertainties remain. The status of two species are somewhat unclear (S. marginata and S. tolteca), calling for taxonomic evaluation. Furthermore, proposed within-genus groupings deserve examination, as do possible subdivisions within some species (e.g., S. limbata, S. parvidentata). Information on basic morphology and biometry remains incomplete for nearly all species. Extreme examples are S. amazonica, S. costalis, and S. paraensis, for which females have not been described. Live animal research on life history, behavior, and ecology is needed for all species, with the possible exceptions of S. carolina and S. limbata. By reconciling species assignments and consolidating biological information for the 23 consensus taxa, this synoptic review promises to guide subsequent systematic and phylogenetic investigations of the genus Stagmomantis.


Fig. 2. Pronotum length (millimeters) for newly emerged adult females in the Þeld. Date of adult emergence is on x-axis; numerical dates are Julian dates, where 1 August ϭ 213. (a) 2008, n ϭ 27 females. (b) 2009, n ϭ 37 females. (c) 2010, n ϭ 20 females. (d) 2011, n ϭ 29 females. 
Fig. 3. Predation events by adult females in the Þeld, 2008Ð2011. (a) Prey length (centimeters) by year. Sample sizes are given in parentheses; the Þrst number represents all prey, except conspeciÞc males; the second number is the number of conspeciÞc males. Black diamonds: mean ( Ϯ SE) prey length, excluding conspeciÞc males. White diamonds: mean ( Ϯ SE) prey length, including conspeciÞc males. (b) Female pronotum length (millimeters) and prey length (centimeters). White squares: conspeciÞc males ( n ϭ 10); black diamonds: all other prey ( n ϭ 63). Vertical dashed lines divide the 63 nonmale predation events into thirds, sorted by female pronotum length. 
Fig. 4. Date of Þrst measurement of dorsoventral abdominal girth (millimeters) for newly emerged adult females in the Þeld and subsequent rate of abdominal expansion (millimeters per day ), 2008Ð2011. Numerical dates are Julian dates, where 1 August ϭ 213. Crosses: 2008; black circles: 2009; triangles: 2010; white circles: 2011. Possible outliers are indicated by the letters A, B, and C. 
Fig. 6. Female pronotum length (millimeters) and measures of reproduction. (a) Fecundity (number of mature eggs in body at death). White circles: 2006 ( n ϭ 18). Black circles: 2008 ( n ϭ 12). (b) Mass of Þrst ootheca (g), 2010 ( n ϭ 26). 
Consequences of Intraspecific Variation in Female Body Size in Stagmomantis limbata (Mantodea: Mantidae): Feeding Ecology, Male Attraction, and Egg Production

December 2013

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70 Reads

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12 Citations

Environmental Entomology

Body size is an important feature of organisms, influencing many components of life history and fitness, such as feeding success and reproductive output. Body size is considered especially salient for solitary predators, whose food intake hinges on individual predation success, which in turn is often driven by the relative sizes of predator and prey. The current study examined intraspecific variation in adult female length and its fitness consequences in a solitary predator, the praying mantid Stagmomantis limbata Hahn. Through a 5-yr integration of observational and experimental work in the field and captivity, we investigated the relationship between female pronotum length and prey size, diet breadth, male attraction, and measures of egg production (fecundity and ootheca mass). We found that longer females ate longer prey in the field and showed greater breadth of prey size than shorter females. Longer females did not necessarily feed at higher rates in the field, as measured by the rate of abdominal expansion. Female length failed to show significant effects on male attraction or on the incidence of cannibalism. Longer females had higher fecundity (mature eggs in body at death) and laid heavier oothecae than shorter females. In nature, longer females consistently emerged as adults earlier in the season than shorter females. Shorter female adults emerged when feeding rates were higher in the field, suggesting an incidental ecological benefit of shorter adult size.


Table 1 Results of maleemale fights in nursery-web spiders
Time that males with and without a nuptial gift took to run across a 1m stick. Numbers in bars are sample sizes.
Gift carrying in the spider Pisaura mirabilis: Nuptial gift contents in nature and effects on male running speed and fighting success

June 2012

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141 Reads

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23 Citations

Animal Behaviour

Males of the spider Pisaura mirabilis offer prey items as nuptial gifts to females. While gift giving in this species has received attention in captivity, the ecological context of these gifts remains largely unknown. First, we examine the occurrence of gift carrying by males in nature. Field data reveal that gift-carrying males are frequent in nature (40% of captured males), and that all gifts contain fresh arthropod prey. Gift mass was positively correlated with the longest diameter of the gift. Thus, males do not appear to 'cheat' by inflating their gifts with inedible items, air or loosely wrapped silk. Second, we examine two ecological costs to gift-carrying males: the effects of gifts on male running speed and male fighting success. Gift carrying reduced male running speed, but did not affect male fighting success in male-male contests. The former result is the first demonstration of a transportation cost associated with gift carrying in an arthropod. (c) 2012 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.


Sexual Conflict Over Spermatophore Attachment in a Nuptially Feeding Cricket

June 2011

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67 Reads

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3 Citations

Ethology

Sexual conflict is implicated in the evolution of nuptial feeding. One function of nuptial gifts lies in mating effort, where the female’s eating of the gift reduces her likelihood of prematurely terminating sperm transfer. We test several ideas regarding sexual conflict in the nuptially feeding wood cricket Nemobius sylvestris. In this cricket, males pass two kinds of spermatophores to females: spermless microspermatophores and larger sperm-filled macrospermatophores. Females may palpate the males’ forewing secretions as a possible additional form of nuptial feeding. We manipulated male mobility and female feeding regime to examine the effects on spermatophore transfer, macrospermatophore attachment duration, and palpations of the males’ forewings. Neither male confinement nor female feeding regime affected the occurrence of macrospermatophore transfer. Males transferred the macrospermatophore sooner to low-food females than to high-food females. Males that were freely mobile (unconfined and sham treatments) had longer macrospermatophore attachment durations than confined males, while female feeding regime did not affect attachment duration. The overall occurrence of female palpation was not significantly affected by female feeding regime. However, high-food females were more likely to perform short palpations before microspermatophore transfer, while low-food females were more likely to palpate after macrospermatophore transfer. Sexual conflict is evident in that males appear to guard against premature removal of the sperm-filled macrospermatophore. Low-food females appear to facilitate the early transfer of the macrospermatophore, while being more likely to perform relatively long post-insemination palpations than better-fed females.


Eggs-per-recruit model for management of the California market squid (Loligo opalescens) fishery

April 2011

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85 Reads

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10 Citations

We develop a per-recruit model for the management of the California market squid (Loligo opalescens) fishery. Based on recent confirmation of determinate fecundity in this species, we describe how catch fecundity (i.e., eggs remaining in the reproductive tracts of harvested females) can be used to simultaneously infer fishing mortality rate along with management reference points such as yield-per-recruit, spawned eggs-per-recruit, and proportional egg escapement. Rates of mortality and egg laying have important effects on these reference points. Somewhat surprisingly, increasing the rate of natural mortality decreased spawned eggs-per-recruit while increasing proportional egg escapement. Increasing the rate of egg laying increased both spawned eggs-per-recruit and egg escapement. Other parameters, such as the maturation rate and gear vulnerability of immature females, affected the reference points. In actual practice, the influence of these parameters for immature squid may go undetected if immature squid are excluded from analysis of the catch. Application of this model to routine management is feasible but requires refinement of sampling procedures, biological assumptions, and model parameters. This model is useful because it is grounded on empirical data collected relatively inexpensively from catch samples (catch fecundity) while allowing for the simultaneous calculation of instantaneous fishing mortality rate and egg escapement.


Behavioral and morphological variation within the Mantodea

December 2010

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26 Reads

Recent behavioral and phylogenetic studies have revived appreciation for variation in behavior and morphology in the Mantodea, both within and between species. In the bordered mantid Stagmomantis limbata, variation in the occurrence of sexual cannibalism is influenced by ecological factors, such as female feeding regime. Female morphology, such as prontoum length, is affected by the environment, with pronotum length decreasing with the date of adult emergence in field populations. This variability in female length within a season produces a broad range of degree of sexual dimorphism (male length/female length x 100%), from 50% to over 90%. Interspecific variation in behavior (sexual cannibalism) and morphology (body length), while based on a somewhat nascent dataset, is considered in a phylogenetic framework.


Figure 3: Fecundity (number of mature eggs in body at death) for females on High and Low diets, 2005–2008. Tests are Mann–Whitney U-tests, where adjusted α = 0.05/4 = 0.013, NS: P > 0.013, *P < 0.013. (a) 2005. Mann–Whitney: U = 88, n = 8 (High), n = 11 (Low), P < 0.001. (b) 2006. Mann–Whitney: U = 234, n = 18 (High), n = 13 (Low), P < 0.001. (c) 2007. Mann–Whitney: U = 25, n = 5 (High), n = 5 (Low), P < 0.01. (d) 2008. Mann–Whitney: U = 88, n = 12 (High), n = 9 (Low), P < 0.001.
Effects of female feeding regime in a sexually cannibalistic mantid: Fecundity, cannibalism, and male response in Stagmomantis limbata (Mantodea)

October 2010

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393 Reads

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51 Citations

Ecological Entomology

1. Female feeding regime exhibited a cascade of effects on reproductive biology and behaviour in the mantid Stagmomantis limbata (Hahn). 2. Well-fed females (High diet) consistently attained greater body mass, thicker abdomens, and higher fecundity than food-limited females (Low diet) in four annual generations. Dorso-ventral abdominal thickness strongly correlated with fecundity. 3. In an experiment in captivity, Low diet females were more likely to cannibalize males than High diet females. 4. High diet females attracted more males than Low diet females in several contexts. In terms of long-range attraction, caged High diet females attracted more males in the field. In terms of short-range attraction, males preferentially mounted and copulated with High diet females in captive paired choice trials. In naturally-occurring pairs in the field, a preference for females with thick abdomens was evident. These results point to two possible explanations: male choice for well-fed females and state-dependent female pheromone emission. 5. Two years of field data indicate considerable variation in female feeding success, as measured by abdominal thickness. Average feeding success in nature appears to be relatively high, when comparing the rate of abdominal expansion in the field to captive females maintained on High diets. 6. In the field, nearly all mounts and copulations occurred in September in both years, when female feeding success is expected to be high. If mating activity occurs during a window of high female feeding success, so as to reduce the risk of cannibalism for males, then the female feeding regime may be implicated in the phenology of mating activity in cannibalistic species.


Citations (12)


... After the production of the egg sac, the female does not feed and carry over it for about three weeks (Austad & Thornhill 1986, Stålhandske 2001a. Wrapping gift with the silk prolongs copulation duration (Lang 1996, Maxwell & Prokop 2018, entices acceptance of the gift (Beyer et al. 2021) and increases male control over nuptial gift as a prevention against its stealing by the female (Andersen et al. 2008). Stålhandske (2002) found that gifts that were painted extra-white with watercolour were accepted more readily by females than unmanipulated gifts or gifts painted brown. ...

Reference:

No Evidence for the Sensory Trap Hypothesis during Courtship in the Gift-Giving Spider Pisaura mirabilis (Clerck, 1757) (Araneae: Pisauridae)
Fitness effects of nuptial gifts in the spider Pisaura mirabilis: Examination under an alternative feeding regime

Journal of Arachnology

... Sexualkannibalismus wurde nun einerseits als adaptives (Buskirk et al. 1984, Birkhead et al. 1988, Prenter et al. 2006) und in anderen Fällen hingegen als nicht-adaptives Verhalten gedeutet (Henriksson 1997). Prokop & Maxwell (2016) konnten jedoch zeigen, dass die als nicht-adaptiv geltende Verwechslungshypothese (Elgar 1992), bei der davon ausgegangen wird, dass der Kannibalisierung eines Geschlechtspartners eine Verwechslung mit einem Beutetier zugrunde liegt, zumindest für M. religiosa nicht zutreffend zu sein scheint. Daher ist davon auszugehen, dass größtenteils adaptive Komponenten zum Auftreten von Sexualkannibalismus beitragen, die insbesondere als für Weibchen vorteilhaft angesehen werden können. ...

Female predatory response to conspecific males and heterospecific prey in the praying mantis Mantis religiosa: evidence for discrimination of conspecific males

Journal of Ethology

... This aligns with developmental patterns in Hierodula species (Leong 2009;Raut et al. 2014;Mirzaee et al. 2022a). Similar variations in the number of moults were observed in Stagmomantis limbata by Maxwell (2014a), proposing it as a "bet-hedging" strategy for developmental diversity (Maxwell, 2014b). Such strategies can enhance the survival of mantid populations, especially in unpredictable environments. ...

Developmental Patterns in Stagmomantis limbata (Mantodea: Mantidae): Variation in Instar Number, Growth, and Body Size
  • Citing Article
  • July 2014

Journal of Orthoptera Research

... In this study, we examine coloration, color change, camouflage strategies, and adult movement in a praying mantis, Stagmomantis limbata (Hahn), a species native to western North America and Central America (Maxwell 2014). Stagmomantis limbata consumes diverse insect prey and is itself subject to predation by spiders, birds, insectivorous mammals, and conspecifics (Maxwell and Frinchaboy 2014). ...

A synoptic review of the genus Stagmomantis (Mantodea: Mantidae)
  • Citing Article
  • February 2014

Zootaxa

... In this study, we examine coloration, color change, camouflage strategies, and adult movement in a praying mantis, Stagmomantis limbata (Hahn), a species native to western North America and Central America (Maxwell 2014). Stagmomantis limbata consumes diverse insect prey and is itself subject to predation by spiders, birds, insectivorous mammals, and conspecifics (Maxwell and Frinchaboy 2014). Body color in S. limbata is variable, with individuals ranging continuously from shades of green to brown, and some individuals exhibiting multiple colors (Roberts 1937;Maxwell 2014). ...

Consequences of Intraspecific Variation in Female Body Size in Stagmomantis limbata (Mantodea: Mantidae): Feeding Ecology, Male Attraction, and Egg Production

Environmental Entomology

... Our measurements showed that the mean female consumption rate in egg sacs was 0.016 g. This value falls well within the range of weights of nuptial gifts found in the study area (Prokop & Maxwell 2012, Prokop & Semelbauer 2017. It suggests that males of P. mirabilis optimise gift weight in order to satiate female feeding requirements. ...

Gift carrying in the spider Pisaura mirabilis: Nuptial gift contents in nature and effects on male running speed and fighting success

Animal Behaviour

... Batsavage 2009) and was modeled as 0 for age 0, 0.3 for age 1, and 1.0 for ages 2+. Our simulations assumed that F remained constant over time, which is common when projecting fishery status by using deterministic biological reference points (e.g., spawning potential ratio [SPR]) to identify sustainable harvest rates (Maxwell et al. 2005). ...

Eggs-per-recruit model for management of the California market squid (Loligo opalescens) fishery
  • Citing Article
  • April 2011

... Adult females have reduced wings and are flightless, whereas males have longer wings and fly in nature (Maxwell and Frinchaboy 2014). Also, the mating system of S. limbata involves sedentary females releasing sex pheromones and males moving to find females, generally through a combination of longer distance flight, followed by shorter distance approach via crawling (Maxwell 1999;Maxwell, Barry, and Johns 2010;Maxwell, Gallego, and Barry 2010). ...

Examinations of Female Pheromone use in Two Praying Mantids, Stagmomantis limbata and Tenodera Aridifolia Sinensis (Mantodea: Mantidae)

Annals of the Entomological Society of America

... Adult females have reduced wings and are flightless, whereas males have longer wings and fly in nature (Maxwell and Frinchaboy 2014). Also, the mating system of S. limbata involves sedentary females releasing sex pheromones and males moving to find females, generally through a combination of longer distance flight, followed by shorter distance approach via crawling (Maxwell 1999;Maxwell, Barry, and Johns 2010;Maxwell, Gallego, and Barry 2010). ...

Effects of female feeding regime in a sexually cannibalistic mantid: Fecundity, cannibalism, and male response in Stagmomantis limbata (Mantodea)

Ecological Entomology

... It is a species closely related to the genus Petaloptila. Males of the species Nemobius sylvestris (Bosc, 1792) secrete a substance on the dorsum of their right tegmina that is actively palpated by the female during copulation [78,83]. Nemobius interstitialis Barranco, Gilgado, and Ortuño, 2013 presents a total loss of stridulatory capacity, and its tegmina have a proliferation of secretory pores much higher than its congenera [84]. ...

Interactions Between Multiple Forms of Nuptial Feeding in the Wood Cricket Nemobius sylvestris (Bosc): Dual Spermatophores and Male Forewings
  • Citing Article
  • December 2008

Ethology