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Raising Neoconocephalus triops (Orthoptera: Tettigoniidae) from Egg to Adult




Introduction

My love for this species began with one very special individual. The first Neoconocephalus triops that I met was Nehemiah, an overwintering adult male who flew to my door on November 12, 2020. I adopted him that evening and he lived with me until his death in the late morning hours on August 29, 2021. Nehemiah sang magnificently, with his first song on February 24, 2021, and his final song on July 21, 2021, directed to his special lady whom I had added to his cage. Here's a VIDEO of Nehemiah singing (with other background singers) on June 25, 2021.
It was Nehemiah who caused me to fall in love with this species and to adopt and care for many others after him:



This article is dedicated with heartfelt gratitude to the human and to the katydid who made my study possible: Dr. Oliver Beckers, who has been my exceedingly patient and extraordinarily kindhearted mentor as I encountered and raised my very first Neoconocephalus katydids, and Nehemiah, the katydid who taught me to love his species.


The "triops" in the name Neoconocephalus triops ("three eyes") refers to the curved dark marking on the ventral side of the fastigium (cone) that gives the appearance of a third eye.


Ventral view of fastigium:


A diagnostic feature of this species is a fastigium that is wider than it is long:


A note about classification: The Singing Insects of North America (SINA) website and a few others list Neoconocephalus in the subfamily Copiphorinae, while BugGuide and others list the coneheads under subfamily Conocephalinae. I believe the latter is the newer classification.



This article includes the following sections: 1) Finding the Katydids, 2) Housing and Care, 3) Life Cycle, Moulting, Overwintering and Diapause, 4) Communication and Song, 5) Mating, 6) Laying Eggs, 7) Parasitism, 8) Geriatrics, 9) Observations, and 10) Questions and Resources, followed by References and Acknowledgements.



Finding the Katydids

While Neoconocephalus triops can be abundant in some areas, in my particular area, they are uncommon and appear to be quite hyperlocalised. I found a single spot in this part of central Texas where N. triops were abundant, a grassy area near a pond. And even with this, the species was completely absent in 2022. I had been able to collect an initial supply of adults in Summer 2021 and raised an F1 generation that overwintered and then an F2 generation in Summer 2022. However, following the pattern of the N. triops in the wild, not a single egg hatched for an F3 generation. Just as in the wild, they simply died out. So I am writing from what I learned through raising an F1 and F2 generation of Neoconocephalus triops.

In central Texas, Neoconocephalus triops are bivoltine: spring adults produce a summer generation and summer adults produce fall babies who overwinter as adults and become sexually mature in very early spring. (Note: a smaller number of spring-born nymphs become adults in summer but do not sexually mature: instead, they diapause through winter and fully mature the following spring.) The best place to look for this species is in habitats with tall native grasses, preferably near a water source. It's easiest to find inhabited locations by searching at night by sound. If there are plenty of males singing, you can guess there will also be females around feeding on grass. I have found that the two hours after complete darkness (after astronomical sunset) is the best time to search for this species. You can also search during the day at the bases of clumps of grass.

This is the best habitat I have found for Neoconocephalus: tall native grasses near a water source:


I use 4"x4" clear plastic containers with lids to catch the insects without harming them. I approach very slowly and simply close in on the insect. Singing males and feeding females can be easy to catch in areas where they are abundant. In areas where they are sparse, I've observed, they tend to be much more skittish and will quickly fly away from an approaching human.



Housing and Care

CARING FOR ADULTS

For housing N. triops, I use large mesh butterfly cages measuring 36 inches tall and 24 inches wide. I provide potted plants with a combination of grasses, specifically inland sea oats (Chasmanthium latifolium) along with various native grasses that I pull up and add to pots. In addition, I collect grass seeds from natural areas to feed these katydids.

This is a sample cage for Neoconocephalus adults:


In addition to the native grasses, I keep a continuous supply of small corn plants (Poaceae) for my Neoconocephalus triops. I start corn seed in 4-inch pots and add the plants to the cages as soon as they are at least six inches tall. My katydids very quickly eat the plants up, so I add new starter plants every week.


For supplemental foods, I add the following: organic Romaine lettuce, organic apple and carrot slices, rolled oats, Fluker's cricket powder, tropical fish flakes, pecans/almonds/walnuts, butterfly sponges, and fresh water.

Here are N. triops munching on apple and lettuce:


Here are N. triops munching on oats:


Here are N. triops on a butterfly sponge and drinking water:


VIDEOS:

Here's a VIDEO of a senior adult eating apple.

Here's a VIDEO of N. triops eating oats,

and a little bit closer VIDEO of N. triops eating oats.

Here's a VIDEO of N. triops drinking water.



CARING FOR NYMPHS

I set up similar cages for nymphs as for adults, except that nymph cages must contain fresh pots of wheatgrass, very important for nymphal development.

This is an example of a nymph cage:


Instructions for Preparing Wheatgrass:

At least two weeks before any Neoconocephalus triops eggs hatch, acquire organic wheatgrass seeds from organic hard red wheat and start your grass.
There are various instructions online for growing wheatgrass from seeds. All methods involve soaking the seeds well before planting. Timing recommendations vary with different instructional resources, so I will describe what I do.
After two days of soaking and rinsing, I plant seeds directly into four-inch pots, covering the top of the soil with seed and then adding a very thin layer of potting mix on top.
I put the pots in plastic shoebox containers and then inside a large cardboard box with a single layer of newspaper loosely covering it. I spray the plants each morning and evening for several days. Once the tallest sprouts get to nearly one inch, I remove the containers and allow the plants to receive light.


Give each set of pots you are working with at least two weeks to mature enough to feed to nymphs.


The nymphs will crawl up onto the plants and eat as well as rest on the plants. Be very careful when handling the plants because the nymphs will be well-hidden and well-camouflaged in the wheatgrass. You can very carefully add small amounts of water to the soil, but you cannot mist because misting can harm or kill the delicate nymphs.

When the grass turns brown and you need to replace the plant with a fresh one, carefully inspect every blade of grass for nymphs. When you find a nymph, place the blade of grass from the old plant next to the new plant and allow the nymph to crawl onto the new plant.
Nymph on wheatgrass:


I always keep a "holding cage" for the pots of dead grass that I remove. That is, the used plants remain for several days inside a separate closed cage just in case I have missed a nymph. Young nymphs camouflage quite well, so it's best to be overly cautious when handling the pots of wheatgrass.

In addition to wheatgrass, I provide nymphs with apple slices, organic Romaine lettuce, butterfly sponges, water sponges, rolled oats, and cricket powder.

Note that water and nectar solutions must be soaked into the sponges and not left as a liquid in a dish in order to ensure that no one drowns.

The youngest nymphs are most interested in munching on wheatgrass and butterfly sponges.
Here is a VIDEO of a three-day old nymph on a butterfly sponge.

Here are nymphs on apple, lettuce, cricket powder, and butterfly sponge:



Here's a VIDEO of male and female seventh-instar nymphs munching on oats.



Life Cycle, Moulting, Overwintering and Diapause

As with all Orthoptera, Neoconocephalus triops are hemimetabolous, undergoing incomplete metamorphosis: egg, nymph, adult.

EGGS

For the N. triops that I raise, the time from hatching to adult is five weeks. Eggs are oblong (elongate), cream-coloured, about five millimetres long, and are "stacked" inside grass stems. Eggs generally hatched five weeks or longer after being laid.

Neoconocephalus triops eggs:


This is an egg just before hatching:



NYMPHS

Nymphs go through seven instars before moulting to adult.

Hatching:

Nymphs must work their way out of the egg from inside the stem, which is an arduous task that can easily fail.
This nymph has made it onto a blade of grass and is resting after the exhausting effort of freeing himself from the egg. The body length is about the same size as an egg, around 5 mm, and the antennae are disproportionately long and extremely delicate in first and second instars.


This nymph is one day old and taking his first sips on the butterfly sponge:


In this case, the nymph has failed to emerge from the egg and dies quickly:


First instar:

These first-instar nymphs are exceedingly delicate and will need good nutrition from grass blades and wheatgrass. The most frequent deaths I had through all life stages were in one- and two-day old nymphs. Many emerge from the egg and then fail within the first two days. I do not know the reason for this.

First-instar nymphs:


Here is a VIDEO of a bit of movement of first-instar nymphs with singing insect sounds in the background.

Second instar:

These are second-instar nymphs, still quite delicate:


Third instar:

By third instar, the nymphs are long and slim:


Fourth instar:

Fourth-instar nymphs start to take shape:


Fifth instar:

In females, the very beginning of an ovipositor can be seen in fifth instar:


Sixth instar:

Note the length of the wing pads and ovipositor in these sixth-instar nymphs:


Seventh instar:

This is a male seventh-instar nymph male crawling over a female seventh-instar nymph:


This female seventh-instar nymph is ready to moult to adult:



ADULTS

This is a teneral adult still darkening:



MOULTING

Green female moulting to adult:


The moulting process in N. triops takes several hours, presenting a very precarious time for the individual. The katydids generally attach themselves to a horizontal surface such as a plant stem or the top of the cage when in captivity and use gravity to assist with pushing out of the exuvia. Failed moultings sometimes occur, usually resulting in death or extreme disfigurement.

After a successful moult, the individual generally eats the exuvia to recover the nitrogenous compounds.
Freshly moulted male eating exuvia:


Here are some close-ups of the exuvia of a male who has moulted from seventh-instar to adult:


Though uncommon, I did observe a very few occasions where the exuvia of a moulting katydid was stolen and eaten by a different individual. Here the male seventh-instar nymph eats the exuvia of a freshly-moulted-to-adult female.


Here is a VIDEO of a few seconds of a female slowly pushing her way out of the exuvia.

Brown female moult to adult series:



This female is moulting from sixth-instar nymph to seventh-instar nymph. Note the size of the ovipositor in the exuvia.


See VIDEO of her freeing her ovipositor from the exuvia.

See VIDEO of her eating the exuvia.

This is a different VIDEO of a female eating the exuvia with night sounds from other singing insects.


OVERWINTERING and DIAPAUSE

In my experience with my F1 overwintering adults, I had 38% (49 of 130) die during diapause. The deaths were pretty evenly spread out over the full winter with more occurring earlier and fewer occurring as spring approached. I have no idea why they died and if something similar or even worse happens in the wild.

Because Texas winters are so capricious, quickly shifting back and forth from Arctic freezes to days above 80°F / 27°C, it seems possible that diapause is continually interrupted (by temperature changes and other natural and man-made disturbances), causing the katydids to deplete fat storage. In captivity (no AC/heat, so conditions were similar to outside but less extreme), my katydids shifted between diapausal and active fairly regularly during the winter, likely due to natural Texas-winter temperature changes. My guess is that an even larger percentage die during diapause in the wild, but I have not found any information on diapause survival in natural conditions. It is known that this tropical katydid species reproduces year-round in the tropics, twice a year in the southern U.S., and once per year farther north (Beckers, 2008). I wondered if winter diapause is a relatively new evolutionary adaptation for this species. Dr. Beckers suggests that since this tropical species is somewhat new to the colder temperatures of the southern U.S., selection may be favouring the more winter-hardy individuals. "Hence, what seems to be a huge loss is just them evolving from a tropical (non-diapause) lifestyle to a more temperate lifestyle" (Dr. Oliver Beckers, personal communication, May 2022). Dr. Beckers further explains that Neoconocephalus triops enter into reproductive diapause rather than conventional diapause (personal communication, March 2022). Perhaps this "partial diapause" makes it even more difficult to conserve energy through the cold winter months.



Communication and Song

My spring-born N. triops generally begin singing about one week after moulting to full adult, and my overwintering adults begin singing in February.
Among my F1 overwintering generation, the first true songs occurred on February 7, 2022, 8:20pm CST at 11.1°C / 52°F and then again on 02.10.2022, 7pm and 7:16pm - 7:25pm at 13.9°C / 57°F. (Before "true singing," there were small buzz noises at times, but no full song.) Singing continued after that any time indoor temperatures were around 13°C or above at 7pm, which was on and off, as intermittent hard freezes continued through mid-March.

QUIVER COMMUNICATIONS

I became aware of the fact that my katydids, both male and female, were communicating through "quivering" without any human-audible sound associated with it. I discovered that this quivering or tremulation has been well-researched.

As an example of katydid tremulation, A. K. Morris et al. describe vibrational communication in Myopophyllum speciosum (Orthoptera: Tettigoniidae) stating "Pairing is completed with vibrational signals, generated at closer range by body oscillation (tremulation). Two distinctive vibrational motor patterns, short and long, are produced by both sexes" (Morris et al., 1994).
Citing various sources, Hannah M. ter Hofstede et al. state, "In addition to acoustic signals, many katydid species in the subfamilies Conocephalinae and Pseudophyllinae produce vibrational signals that travel through plants" (Hofstede et al., 2020). Regarding Pseudophyllinae, Inga Giepel et al. explore vibrational communication with respect to predation risks by bats: "Male Neotropical pseudophylline katydids produce both acoustic and vibrational signals (tremulations). Females reply to male signals with tremulations of their own, and both sexes walk to find one another" (Geipel et al., 2020).

Preflight warm-up vs. quiver communications:

Preflight and pre-hop warm-up differs from quiver communication with the former being in-place and not directed to others.
As an example of pre-hop warm-up, this VIDEO shows a female warming up in order to hop.

From what I could see of quiver communications, it occurs in both sexes, but more frequently in males, and is directed toward others.

Here are a few VIDEO examples:

Interestingly, this pair has just completed mating, but the male continues to communicate with the female using quivering. The sounds are from off-camera individuals in different cages. (This pair had their own cage, allowing me to observe them more closely.)
VIDEO of male communicating with female after mating.

This is the same pair. Note that the female does not have to be looking at the male during quiver communication.
VIDEO of male communicating with female after mating.

VIDEO of male using quiver communication.

The male in this VIDEO is directly communicating with the nearby female without human-audible sound.

In this VIDEO, males and females are communicating using quivering, but the actual singing is from different off-camera individuals.

Regarding the actual song, Dr. Oliver Beckers told me "... they definitely have some song components in the ultrasound range. A colleague of mine did an experiment and found that adding the ultrasound component to an artificial call increases the accuracy of the approach to the call. Note that ultrasound deteriorates much faster over distance than lower frequencies, i.e., they are only audible to the female at closer range" (personal communication, July 2022). I wonder if quiver communication involves not only vibrations but possibly an ultrasound component.

SONG

The Neoconocephalus triops song can be either a continuous trill or a "versed" trill regularly interrupted at short intervals.

Of all the katydids and crickets I've observed singing, Neoconocephalus triops appear to me to expend the greatest amount of energy to stridulate. Singing appears to involve the whole body, often with quite jerky movements of the abdomen as if to assist in stimulating the thoracic muscles. My impression is that the vigorous stridulation of N. triops wears on males, causing their wings to become tattered and contributing to their earlier deaths when compared to females. That is, singing appears to be disproportionately expensive for males, not only in inviting the risk of predation but also in the sheer amount of effort exerted in this costly call for a mate.

I've noticed that males often warm up their muscles before singing: the wings and abdomen are making the motions of singing, but there is no sound. After a few seconds of this, the male begins his audible song.

With my N. triops spread out in several different cages, singing generally begins and pauses around the same times for all cages.

Here are a few VIDEO examples of calling:

Song VIDEO

Song VIDEO

Song VIDEO

Song VIDEO, Sound Only (with other singers)


Male Competition:

Here's a VIDEO of male competition for the Katydid's Got Talent competition.

And … a bit of a Boxing Match VIDEO for Katydid Fight Night.


Mating

I haven't seen Neoconocephalus mating in the wild, but I was able to observe mating within my captive N. triops community.

In one of the best katydid articles I've ever encountered, Dr. Karim Vahed describes male cercal morphology in relation to brief versus prolonged copulation. The species most similar to N. triops described in his article is Ruspolia nitidula (Conocephalinae) with the grasping cerci associated with prolonged copulation.

While I have seen and photographed mating, I have never gotten to see the initiation of the process. I've only seen actual mating and separation. The maximum amount of time I have seen N. triops remain in copula (from some time after the beginning until separation) was 40 minutes.
Dr. Vahed (2014) states, " …longer copulation following spermatophore transfer was associated with larger ejaculates, across species with reduced nuptial gifts. Our results demonstrate that nuptial gifts and the use of grasping cerci to prolong ejaculate transfer are functionally equivalent." From what I could see, there was no obvious spermatophylax as is clearly present in, for example, Amblycorypha huasteca and A. uhleri. Likely the prolonged copulation ensures the optimal transfer of spermatozoa.

Images of male grasping cerci:


Generally while mating, the pair remain in one place, but I do occasionally see the female leading them to a different location.

Images of Neoconocephalus triops mating:


Here's a VIDEO of mating.

It appears that the female eats the ampulla after mating.


Here's a VIDEO of a female eating the ampulla after mating.


Laying Eggs

A female lays eggs by first chewing a "pilot hole" or starter hole into a relatively thick part of a grass stem. She then gradually works her ovipositor downward into the stem and (apparently) releases several eggs stacked vertically along the path as she pulls out.

Female chewing starter hole in grass stem:


Females laying eggs:


Here is a longish VIDEO of female laying eggs with night singing sounds.

And another VIDEO of a female laying eggs in the base of the grass stems.

Here is a VIDEO of a female finishing preparing the stem and then laying eggs.


Parasitism

One of the major predators of Neoconocephalus triops in my area is the parasitoid fly Ormia lineifrons (Diptera: Tachinidae).
Ormia lineifrons adult female and puparia:


The expert on this and other fly predation of Neoconocephalus is Dr. Oliver Beckers of Murray State University, Kentucky, along with Dr. James O'Hara of the Canadian National Collection of Insects.

In my experience with rearing N. triops, I tried hard to avoid encountering this fly, but since I collected my initial adults from the wild, I did indeed learn rather quickly about fly predation.

Ormia lineifrons females lay eggs near the host, and the larvae, which hatch almost immediately, burrow into the host and develop inside. Fully developed larvae exit the host, killing the host in the process. Larvae pupate almost immediately upon exit, generally remaining in the pupal stage for around 12-13 days before emerging as adults (Dr. Oliver Beckers and Dr. James E. O'Hara, personal communications, August 2021).

Here, the mature larva of an Ormia lineifrons has exited a male Neoconocephalus triops to pupate. The katydid has been killed by the parasitoid fly.


Dr. Oliver Beckers presents fascinating detailed studies of these animal interactions in a series of elegant papers and adds an examination of superparasitism of N. triops as well as parasitism by an undescribed species of a Neomintho genus Tachinid fly (Beckers, 2022).

For rearing purposes, it is safest to isolate newly collected males for a couple of weeks before mixing them with others in captivity. The fly can also be controlled by immediately and regularly ridding the cages of any puparia or adults who might emerge.


Geriatrics

Neoconocephalus triops are fairly long-lived katydids in captivity and many live for more than a year after hatching from the egg.

Because my katydids are kept in fairly ideal conditions, they live out their full natural lives and grow old inside their habitats. As adult Neoconocephalus triops age, they show signs of ageing similar to other katydids. They may begin to drop limbs, usually beginning with hind legs. Tarsi and tibiae may be dropped. Antennal parts break off. The abdomens may swell, especially in females, and overall colouration may change, either darkening or fading. Wings become tattered and may tear.

Two senior males and one senior female showing signs of ageing:


At this stage, the katydids become less alert and less mobile. I provide them with easy access to foods: they are especially interested in butterfly sponges (nectaring) and apple slices. Lettuce, rolled oats, and wheatgrass are also desired treats among seniors.

In this VIDEO two very senior adults climb up to munch on wheatgrass (with night singing sounds).

Here is a VIDEO of two senior citizens munching on an apple with night singing sounds.

Here is a VIDEO of a senior adult on a butterfly sponge with night singing sounds.

For all of the creatures I raise through multiple generations, I keep assisted living cages and hospice cages for senior adults in order to keep them comfortable and ensure that they have easy access to nutrition. For N. triops, I add sticks to allow the katydids to climb up to their preferred perching plants, and I ensure that all food and drink is easily accessible.

Sample Assisted Living cage:



Observations

Winter vs. Summer size:

One thing that I anecdotally observed is that my winter adults tended to be smaller than summer adults. Winter adults need a higher build-up of fat to survive diapause, but perhaps higher fat in a smaller body is the optimal combination for an adult who must diapause through winter.

As an example, these two green female specimens show winter (55mm) vs. summer (60mm) size.


Heavy Sleepers:

My different species of insects appear to experience very different levels of sleep. Neoconocephalus triops is a species that enjoys a deep sleep. They are slow to awaken and quite lethargic when first awakened. Their most active period is the few hours after total darkness.
Just a random personal thought: In my area, this species overwinters as adults. As such, they have a built-in adult diapause mechanism to allow them to survive the winter with little to no nutrition. Perhaps that mechanism also applies to daily sleep, allowing the insects to enter into a more torpid state than do other species.

Side Sleeping:

One interesting behaviour in this species that quite surprised me the first time I saw it is "side sleeping." I often find these guys tilted at around a 45-degree angle to one side while sleeping. It's difficult to get an actual photograph of side sleeping because at the slightest disturbance, such as at the opening of the cage, they straighten up, but here are the best shots of partial side sleeping I could get:


Colour Dimorphism: Green, Brown, and … Pink?

Neoconocephalus triops can be brown or green. In my area, there are more brown katydids in the overwintering generation and more green katydids in the summer generation. There are fewer green katydids who overwinter and most of those are female. I believe the colour dimorphism varies with location and conditions. I would like to have worked more with colour variation, but I failed to get an F3 generation to continue the experiments I had begun. Photoperiod clearly plays a significant role in colour. Dr. J. J. Whitesell did extensive experiments with photoperiod to effect changes in colour, song, fat build-up, and more (Whitesell, 1974). Even so, I wonder if there are other factors that contribute to the colour of individual Neoconocephalus triops, if genetics has any role, and also if colour itself affects other things, such as predation (likely), mate attraction, health, or anything else.

I did not get to follow up with experiments, but I did document my F2 generation with respect to colour phase as follows.

F2 summer adult offspring from F1 overwintering parents – numbers by colour:

Total F2 offspring: 26 green, 44 brown = 70
26/70= 37% green
44/70 = 63% brown

Green & Brown mixed parental cages (The F1 parents of these F2 offspring were in two cages of green and brown mixed N. triops):
female - 12 green, 11 brown
male - 5 green, 8 brown
total 17 green, 19 brown = 36
17/36 = 47% green
19/36 = 53% brown

Brown-only parental cages (The F1 parents of these F2 offspring were in two cages of brown-only N. triops):
female - 6 green, 14 brown
male - 3 green, 11 brown
total: 9 green, 25 brown = 34
9/34 = 26% green
25/34 = 74% brown

Although I was unable to continue my experiments, I got the impression that two brown parents were more likely to produce brown offspring even while seasonality (photoperiod) continued to play the major role.

Pink?

I am unaware of actual erythrism in Neoconocephalus, but I did get a handful of pinkish individuals. They were brown form, both male and female, with distinctive hints of pink.

Pink-tinted Neoconocephalus triops:


After they died, the pink showed up more prominently whereas no such pigmentation occurred after death in my "normal" brown specimens.
Here you can see a pink-tinted individual after death and a comparison with a normal brown individual:


So while there was no actual erythrism, there was a bit of "pinkmentation."


Questions and Resources

QUESTIONS

Just a Few of My Many, Many Questions:

– What other factors besides photoperiod contribute to the colour of individual N. triops? Does genetics have any role? Does colour itself affect other things besides likelihood of predation by visually-oriented predators, such as attractiveness to a mate, overall health, or other?

– How is mating initiated? I assume the female approaches a suitable male, but what transpires then and how does he initially grasp her?

– Do females mate more than once? Based on what I saw after setting up several cages containing only one N. triops pair each, I got the impression (but have no evidence) that they mate more than once.

– Is there any reproductive value to eating the ampulla as the female appears to do after mating? Regarding consumption of a full spermatophylax, apparently absent in this case, many studies have been conducted that show at least some reproductive benefit to the progeny (Gwynne, 1988) and nutritional benefit to the female (Gwynne et al., 1984), and I assume that the ampulla itself at least provides some nutrition to the female.

– For how long can a female store viable sperm?

– How sensitive are these guys to inbreeding?

– Is the evolutionary choice of prolonged copulation vs. spermatophylax production for the purpose of resource conservation / longevity and does it affect fecundity? Is the quality of spermatozoa transfer comparable?

– How much energy is expended in singing and how does it affect the male's longevity? Likewise, how much energy is expended in producing spermatophores, and how does it affect the male's longevity?

– What happens during "quiver communication" or tremulation and what is being communicated? Is it purely vibrational or is there an ultrasound component?

– Why do so many one-day-old and two-day-old nymphs die, relative to all other life stages? (My guess is that they completely exhaust themselves and deplete all their resources during the arduous process of working their way out of the egg and out of the stem.)

– When eggs are stacked together in a single stem hole, does only one nymph generally survive or do a few survive?

– Why did this species evolve to overwinter as adults? What benefit is it to the species to overwinter/diapause as adults rather than as eggs like other Neoconocephalus species?

– What happens during diapause and what is the survival rate in the wild? Is diapause for bivoltine overwintering N. triops comparable to diapause for univoltine overwintering N. triops and are they similarly successful?

– To what degree are these guys migratory? How far do they travel and where/when does this occur? I believe that fall adults migrate to their overwintering grounds and then migrate again in early spring to feeding/breeding grounds. What triggers migration and what informs the routes they take?

– Why were Neoconocephalus triops absent in my area in 2022? (Bigger pictures: why are particular Tettigoniidae species present one year and absent the next?)

– Why did I get no F3, even though conditions were exactly the same as for my very successful F1 and F2 generations?

– a little more twilight zone'ish: Is it possible that N. triops would ever overwinter as an egg?

(Editing Note: this article is still in progress – I will be adding/editing.)



RESOURCES

A few of the articles from the Neoconocephalus guru, Dr. Oliver Beckers:

Beckers, O. M. (2008, July).The Evolutionary Significance of Developmental Plasticity in the Communication System of Neoconocephalus Triops (Orthoptera: Tettigoniidae). https://pdfs.semanticscholar.org/bf5a/3892c1db53d43ec3bc59291011f12f10ef99.pdf

Beckers, O. M. (2022, June 10). Parasitism of the Katydid Neoconocephalus Triops (Orthoptera: Tettigoniidae) by the Tachinid flies Ormia lineifrons and Neomintho sp. (Diptera: Tachinidae). BioOne Complete. https://bioone.org/journals/florida-entomologist/volume-105/issue-2/024.105.0205/Parasitism-of-the-Katydid-iNeoconocephalus-triops-i-Orthoptera--Tettigoniidae/10.1653/024.105.0205.full

Beckers, O. M., & Schul, J. (2008, February 26). Developmental plasticity of mating calls enables acoustic communication in diverse environments. Proceedings of the Royal Society B: Biological Sciences. https://royalsocietypublishing.org/doi/full/10.1098/rspb.2007.1765

Beckers, O. M., & Schul, J. (1970, January 1). Female adaptation to developmental plasticity in male calling behavior. AGRIS. https://agris.fao.org/agris-search/search.do?recordID=US201301867967

Bush, S. L., Beckers, O. M., & Schul, J. (2009, March 1). A complex mechanism of call recognition in the Katydid Neoconocephalus affinis (Orthoptera: Tettigoniidae). The Company of Biologists. https://journals.biologists.com/jeb/article/212/5/648/18971/A-complex-mechanism-of-call-recognition-in-the

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More resources:

Doubell, M., Grant, P. B. C., Esterhuizen, N., Bazelet, C. S., Addison, P., & Terblanche, J. S. (2017, December). The metabolic costs of sexual signalling in the chirping katydid Plangia graminea (Serville) (Orthoptera: Tettigoniidae) are context dependent: cumulative costs add up fast. https://journals.biologists.com/jeb/article/220/23/4440/33679/The-metabolic-costs-of-sexual-signalling-in-the

Josephson, R. K. (1985, July 1). Mechanical power output of a tettigoniid wing muscle during singing and flight. The Company of Biologists. https://journals.biologists.com/jeb/article/117/1/357/5161/The-Mechanical-Power-Output-of-a-Tettigoniid-Wing

Linn, S., & Gillett-Kaufman, J. (2015, December). common name: broad-tipped conehead katydid (suggested common name) scientific name: Neoconocephalus triops (Linnaeus, 1758) (Insecta: Orthoptera: Tettigoniidae: Conocephalinae). Featured Creatures. https://entnemdept.ufl.edu/creatures/MISC/Neoconocephalus_triops.htm

Quinn, M. (2008, November 19). Broad-tipped Conehead. Broad-tipped Conehead - Neoconocephalus triops (Linnaeus 1758). https://www.texasento.net/triops.htm

Schulze, W., & Schul, J. (2001, February 15). Ultrasound avoidance behaviour in the bushcricket tettigonia viridissima (Orthoptera: Tettigoniidae). Journal of Experimental Biology. https://journals.biologists.com/jeb/article/204/4/733/8739/Ultrasound-avoidance-behaviour-in-the-bushcricket

Snyder RL, Frederick-Hudson KH, & Schul J. Molecular phylogenetics of the genus Neoconocephalus (orthoptera, tettigoniidae) and the evolution of temperate life histories. PLoS One. 2009 Sep 25;4(9):e7203. doi: 10.1371/journal.pone.0007203. PMID: 19779617; PMCID: PMC2745656. https://journals.plos.org/plosone/article?id=10.1371/journal.pone.0007203

Strauß , J., Alt, J. A., Ekschmitt , K., Schul, J., & Lakes-Harlan 1, R. (2017, May 2). Evolutionary diversification of the auditory organ sensilla in neoconocephalus katydids (Orthoptera: Tettigoniidae) correlates with acoustic signal diversification over phylogenetic relatedness and life history. Journal of evolutionary biology. https://pubmed.ncbi.nlm.nih.gov/28295793/

Stumpner, A., Dann, A., Schink, M., Gubert, S., & Hugel, S. (2013, January 1). True Katydids (Pseudophyllinae) from Guadeloupe: Acoustic signals and functional considerations of song production. OUP Academic. Retrieved January 8, 2023, from https://academic.oup.com/jinsectscience/article/13/1/157/1754081

Symes, Laurel & Page, Rachel & Hofstede, Hannah. (2016). Effects of acoustic environment on male calling activity and timing in Neotropical forest katydids. Behavioral Ecology and Sociobiology. 70. 10.1007/s00265-016-2157-4.

ter Hofstede, H. M., & Fullard, J. H. (2008, August 1). The neuroethology of song cessation in response to gleaning bat calls in two species of katydids, Neoconocephalus Ensiger and Amblycorypha oblongifolia. Journal of Experimental Biology. https://journals.biologists.com/jeb/article/211/15/2431/17511/The-neuroethology-of-song-cessation-in-response-to

Walker, T. J., & Greenfield, M. D. (1983). Songs and Systematics of Caribbean Neoconocephalus (Orthoptera: Tettigoniidae). Transactions of the American Entomological Society (1890-), 109(4), 357–389. http://www.jstor.org/stable/25078329

and for fun:

Rakesh, M., Aris-Brosou, S., & Xia, X. (2022, June 22). Testing alternative hypotheses on the origin and speciation of Hawaiian katydids - BMC ecology and evolution. BioMed Ecology and Evolution. Retrieved from https://bmcecolevol.biomedcentral.com/articles/10.1186/s12862-022-02037-2


References

Beckers, O. M. (2008, July). The Evolutionary Significance of Developmental Plasticity in the Communication System of Neoconocephalus Triops (Orthoptera: Tettigoniidae). https://pdfs.semanticscholar.org/bf5a/3892c1db53d43ec3bc59291011f12f10ef99.pdf

Beckers, O. M. (2022, June 10). Parasitism of the Katydid Neoconocephalus Triops (Orthoptera: Tettigoniidae) by the Tachinid flies Ormia lineifrons and Neomintho sp. (Diptera: Tachinidae). BioOne Complete. https://bioone.org/journals/florida-entomologist/volume-105/issue-2/024.105.0205/Parasitism-of-the-Katydid-iNeoconocephalus-triops-i-Orthoptera--Tettigoniidae/10.1653/024.105.0205.full

Geipel, I., ter Hofstede, H. M., Page, R. A., Carter, G. G., Litterer, A. S., & Kernan, C. E. (2020, March 16). Predation risks of signalling andsearching: bats prefer moving katydids. Predation risks of signalling and searching: Bats prefer moving katydids. https://royalsocietypublishing.org/doi/epdf/10.1098/rsbl.2019.0837

Gwynne, D. T. (1988, May). Courtship feeding and the fitness of female katydids (Orthoptera: Tettigoniidae). Evolution; international journal of organic evolution. https://pubmed.ncbi.nlm.nih.gov/28564010/

Gwynne, D. T., Bowen, B. J., & Codd, C. G. (1984, January 1). The function of the katydid spermatophore and its role in fecundity and insemination (Orthoptera: Tettigoniidae). Australian Journal of Zoology. https://www.publish.csiro.au/ZO/ZO9840015

Morris, G. K., Mason, A. C., P. Wall, & Belwood, J. J. (1994, May). High ultrasonic and tremulation signals in neotropical katydids (Orthoptera: Tettigoniidae). https://zslpublications.onlinelibrary.wiley.com/doi/abs/10.1111/j.1469-7998.1994.tb05266.x

ter Hofstede, H. M., Symes, L. B., Martinson, S. J., Robillard, T., Faure, P., Madhusudhana, S., & Page, R. A. (2020, April 12). Calling songs of neotropical katydids (Orthoptera: Tettigoniidae) from Panama. Journal of Orthoptera Research. Retrieved January 8, 2023, from https://jor.pensoft.net/article/46371/

Vahed, K., Gilbert, J. D. J., Weissman, D. B., & Barrientos-Lozano, L. (2014, July). Functional equivalence of grasping cerci and nuptial food gifts in promoting ejaculate transfer in katydids. https://pubmed.ncbi.nlm.nih.gov/24724547/

Walker, T. J. (n.d.). Subfamily Copiphorinae (coneheaded katydids). https://orthsoc.org/sina/s160a.htm

Whitesell, J. J. (1974, January 1). Geographic variation and dimorphisms in song, development, and color in a katydid; field and laboratory studies (Tettigoniidae, Orthoptera). https://www.biodiversitylibrary.org/item/90426


Acknowledgements

I owe a tremendous debt of gratitude to the folks who made my study possible:

I want to thank from the bottom of my heart Brandon Woo, Entomologist Extraordinaire, who captured for me the first adults that I used to start my colony. Brandon is as kindhearted and generous as he is brilliant and skillful.

I am deeply grateful to the Neoconocephalus guru, Dr. Oliver Beckers, who has tirelessly answered my questions and mentored me throughout my time in studying this species. It was Dr. Beckers who inspired me and ignited my passion to look more closely at this magnificent species of katydid. He is a deeply compassionate and exceedingly creative and thoughtful scholar whose work will continue to be treasured in the field of entomology. Dr. Beckers generously shared his elegant writings with me, patiently instructed me, pointed me to his fellow experts, and remained a consistently kind and compassionate friend to me whenever I lost some of my beloved katydids or felt discouraged. I am forever grateful to Dr. Oliver Beckers.

I want to thank the delightful and brilliant Dr. Karim Vahed who has graciously answered my many questions and Dr. James O'Hara who patiently shared his expertise to help me with the Ormia lineifrons parasitoid fly.



.

Might grab a few of these whi
Might grab a few of these while I'm at Houston this December.

 
That would be great!
This is an awesome species to raise because the ones here in Texas are bivoltine and overwinter as adults.
They would be harder to find in December, but you might find them at lights (I believe they migrate to overwintering grounds and you might catch some on their way). Or you might get lucky and find some hiding in forested areas. The December ones will be adults but not fully sexually mature yet and, of course, will not be singing.
Please let me know what you find while you are here in December.
I am so glad that you are interested in this species, Hemiptamantis, because it is one of my most beloved species and I think you would be great at rearing these guys.

 
I did hear a few calling in t
I did hear a few calling in the middle of December when I was there a couple years ago.

 
Ah, excellent !!
I look forward to hearing about what you find there. It sounds like they sing longer into the season than do ours here. When I seek them out at night by sound, I look for nearby females perched up high on grasses. I am confident that you will have success and I look forward to reading about your adventures.

Wow, this is a great article!
Wow, this is a great article! I do have a couple notes to add based on my experience with meadow katydids, though
-How is mating initiated? I assume the female approaches a suitable male, but what transpires then and how does he initially grasp her?
In Conocephalus, Orchelimum, and Neoconocephalus, mating is initiated by the male. He lines up alongside the female with his head pointing in the opposite direction, and then lifts up a hind leg and curves his abdomen towards the end of the female's abdomen with the cerci open. Then, after touching the female, he shifts around until the cerci lock in the correct position, and then mating begins. They also do the same quiver/vibration when they approach a female.

– Is there any reproductive value to eating the ampulla as the female appears to do after mating?
In most katydids, the entire spermatophylax structure is a way for the male to provide nutrition to the female, and in many species, it is almost comically large. Producing it is a significant energy expenditure for the male, perhaps even moreso than singing. In all of the katydid matings I've seen, the female never fails to eat the entire thing. I would assume that the additional nutrients are probably beneficial to the female.

– How sensitive are these guys to inbreeding?
Sort of unrelated, but my captive-raised greater angle-wings would simply refuse to inbreed. The males would sing, and the females would respond, but mating never occurred, and I never saw males attempting to mate once coming in contact with a female. Perhaps they know that the females are related, and would not be suitable mates?

Also, how long were your captive adults able to live? I saw the section on geriatrics, but nothing was mentioned about their total lifespans.

Another thing I found interesting was that adult males were rather peaceful with each other. Conocephalus and Orchelimum males will physically fight each other every once in a while, squaring up against each other, attempting to mate, and then trying to bite the limbs/abdomen of the opponent. Towards the end of the season, it is quite common to see males with bite marks on their middle and hind legs. However, a "pecking order" is usually quickly established in captivity, and the losers of fights soon learn to avoid the winners, which avoids future conflict.

The vibrations while singing and before physical activity seems to be something common to Neoconocephalus, but only Neoconocephalus. None of the meadow katydid species do it, but both the N. ensiger and robustus I've seen vibrate vigorously when initiating physical activity, especially at night. I suspect it is probably similar to the vibrations of moths before taking off, perhaps serving to warm up the body when ambient temperatures are low. This would make sense, because Neoconocephalus are more exclusively nocturnal than the other conocephaline genera, and have the large and strong wing muscles needed to generate heat. I recall reading a paper that mentioned N. robustus being able to raise its body temperature by a significant amount, allowing them to continue singing in cold weather. https://www.journals.uchicago.edu/doi/10.2307/1540212

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